UNITED  STATES  DEPARTMENT  OF  AGRICULTURE 

BULLETIN  No.  826 


Contribution  from  the  Bureau  of  Entomology 
L.  O.  HOWARD,  Chief 


Washington,  D.  C. 


PROFESSIONAL  PAPER. 


August  10,  1920 


GENERIC  CLASSIFICATION  OF  THE  HEMIPTEROUS 
FAMILY  APHIDID AE.'  r^S  6 


By  A.  C?  BAKER, 

Entomologist,  Deciduous  Fruit  Insect  Investigations. 


CONTENTS. 


Superfamily  Aphldoidea  

Page. 
2 

Subfamily  III,  Eriosomatinae  

Page. 
62 

Phylogeny  of  the  Aphididae 

3 

Tribe  Eriosomatini  

65 

Key  to  the  subfamilies  of  the  Aphididae.  . 

10 

Tribe  Pemphigini  

68 

Subfamily  I,  Aphidinae 

10 

Tribe  Melphini 

73 

Tribe  T,achnini 

12 

Tribe  Prootphilini 

75 

Tribe  Thelaxini 

20 

Tribe  Fordini  .  . 

77 

Tribe  Callipterini 

21 

Subfamily  IV,  Hormaphidinae  

...  81 

Tribe  Greenideini. 

37 

Tribe  Hormaphidini  

83 

Tribe  Setaphidini 

38 

Tribe  Oregmini  

.   .   .            84 

Tribe  Aphidini 

39 

Tribe  Cerataphidini  

.   .   .            86 

Pnhfamily  TT    Mindarina<> 

61 

Genera  not  placed  

88 

Probably  no  group  of  insects  has  received  more  attention  at  the 
hands  of  economic  entomologists  than  aphids,  or  plant-lice.  Their 
interesting  and  often  complicated  biologies  have  attracted  the  atten- 
tion of  investigators,  not  only  among  entomologists,  but  among 
workers  in  the  larger  fields  of  zoology  and  general  biology.  While 
a  large  amount  of  work  on  the  life  histories  and  biologies  of  aphids 
has  been  done,  corresponding  progress  in  their  classification  has  not 
been  made.  This  is  probably  due  to  several  causes,  su^h  as  the  lack 
of  correlation  of  biologic  and  taxonomic  facts,  and  the  failure  of 
aphidologists  to  consider  sufficiently  the  results  of  the  work  of  others. 

On  account  of  the  great  economic  importance  of  aphids  and  the 
necessity  of  their  study  in  the  development  of  control  measures,  the 
lack  of  knowledge  concerning  their  systematic  relationships  results 
in  much  confusion.  Some  biologic  workers,  in  fact,  do  not  now  at- 
tempt to  give  the  name  of  the  species  being  studied  on  account  of  the 
difficulty  experienced  in  securing  correct  determinations. 


1  This  paper  is  the  first  of  a  series  treating  the  Aphididae.    It  will  be  followed  by  others  dealing  with 
the  economic  importance,  biologies,  and  relationships  of  species  in  the  different  genera. 
141613°— 20— Bull.  826 1 


2  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

The  present  work  was  undertaken  in  the  hope  of  remedying  such 
a  condition  to  some  extent  at  least.  The  genera  of  the  world  have 
been  studied.  Many  workers  have  lent  material  and  the  large  col- 
lections of  the  National  Museum  and  Bureau  of  Entomology  have 
been  drawn  upon.  In  the  National  Museum  collection  a  large  per- 
centage of  types  has  been  available.  To  the  study  of  preserved 
material  have  been  added  embryological,  anatomical,  and  biological 
investigations  that  a  better  understanding  of  the  natural  relationships 
might  be  gained. 

Besides  many  aphidologists  in  this  country  and  abroad,  who  have 
given  helpful  suggestions  and  many  of  whom  have  read  and  criticized 
the  manuscript,  the  writer  is  indebted  to  Dr.  A.  L.  Quaintance,  of 
the  Bureau  of  Entomology,  for  the  f acilities  for  conducting  many  of 
the  biological  investigations  which  to  a  large  extent  have  laid  the 
foundation  for  the  systematic  treatment  here  given. 

Superfamily  APfflDOIDEA. 

There  appear  to  be  two  distinct  families  in  the  superfamily  Aphi- 
doidea.  These  are  the  Aphididae  and  the  Phylloxeridae.  The  present 
paper  deals  only  with  the  Aphididae. 

Members  of  the  Phylloxeridae  differ  markedly  from  forms  belong- 
ing to  the  Aphididae.  In  the  first  place  their  biologies  are  quite 
different  in  that  parthenogenetic  oviparous  forms  occur  during  the 
summer.  In  the  Aphididae  only  the  sexed  females  which  are  pro- 
duced in  the  fall  are  normally  oviparous. 

In  structure  the  two  families  are  separated  at  once  by  the  formation 
of  the  stigma  of  the  forewing.  The  wing  itself  seems  very  little 
different  in  an  Adelges  or  Phylloxera  from  that  in  some  of  the  special- 
ized genera  of  the  Aphididae.  An  examination  of  the  freshly  emerged 
wing,  however,  as  has  been  pointed  out  by  Dr.  Patch,  shows  that  the 
stigma  in  the  Phylloxeridae  is  formed  by  the  radial  sector  and  the 
stigmal  vein  is  the  media.  In  the  Aphididae,  on  the  other  hand, 
the  stigma  is  formed  by  radius  x  and  the  stigmal  vein  is  the  radial 
sector.  The  two  families  may  thus  be  separated  as  follows: 

KEY  TO  THE  FAMILIES  OF  THE  APHIDOIDEA. 

Summer  parthenogenetic  oviparous  forms  produced:    Stigma  formed  by 

the  radial  sector PHYLT-OXERIDAE. 

Only  sexual  oviparous  forms  produced :    Stigma  formed  by  radiusj APHIDIDAE. 

A  word  of  explanation  in  regard  to  the  name  Phylloxeridae  may 
be  necessary.  The  genus  Chennes  was  erected  by  Linnaeus  in  1758 
and  in  1862  was  replaced  by  Psylla  Geoffrey.  For  this  genus  CTiermes 
ficus  L.  was  set  as  type  by  Lamarck  in  1801.  Ficus,  therefore,  be- 
comes ipso  facto  the  type  •  of  Chermes,  and  Chermidae  the  family 
name  of  the  "jumping  plant-lice."  The  family  name  for  the  aphi- 
doidean  group,  therefore,  is  derived  from  the  genus  Phylloxera 
Boyer  (1834). 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  3 

PHYLOGENY  OF  THE  APHIDIDAE. 

In  many  published  classifications  of  the  Aphididae  those  groups 
which  according  to  the  writer's  conception  are  the  most  specialized 
have  been  placed  as  the  most  primitive.  This  is  the  case  with  those 


FIG.  l. — Phytogeny  of  the  hemipterous  fainily  Aphididae. 

insects  forming  in  the  present  classification  the  Eriosomatinae  and 
the  Hormaphidmae. 

A  study  of  the  anatomy  and  the  biology  of  aphids  makes  it  evident 
that  there  are  three  main  groups  of  living  forms  for  which  subfamily 


4  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

names  are  here  used :  Aphidinae,  Eriosomatinae,  and  Hormaphidinae. 
Besides  these  there  is  the  one  species,  in  some  -ways  a  relic  of  the 
past,  forming  th'e  Mindarinae. 

As  will  be  seen  by  the  diagram  (fig.  1),  the  Aphidinae  is  considered 
the  most  primitive  subfamily  of  the  three  main  groups.  This  is 
substantiated  by  the  habits  of  the  insects,  by  the  structure  of  the 
wings,  beak,  etc.,  and  by  the  sexual  forms. 

Practically  all  of  these  insects  are  foliage  or  twig  feeders.  They 
live,  as  a  rule,  in  colonies  and  have  not  developed  any  very  special 
methods  of  We,  such  as  highly  complex  gall  formation.  With  the 
exception  of  the  Mindarinae  the  most  primitive  wing  structure  oc- 
curring in  the  family  is  met  with  here.  The  media  is  most  commonly 
twice  branched  throughout  the  subfamily  and  even  in  the  Mindarinae 
there  is  more  of  a  reduction  than  this.  The  antennae  are  of  six 
segments,  the  largest  number  found  in  the  family,  and  the  sen- 
soria  are  simpler  in  nature  than  those  met  with  in  the  other  two  large 
subfamilies.  The  beak  in  the  Lachnini  shows  also  a  primitive 
condition  in  its  segmentation. 

The  sexual  forms  are  most  primitive  in  the  Aphidinae.  Winged 
sexes  often  occur,  at  least  the  males  are  very  commonly  winged.  Both 
sexes  still  retain  their  beaks  and  feed  on  their  hosts  in  the  same 
way  as  do  the  other  forms,  and  the  ovaries  of  the  female  develop 
normally,  and  she  produces  several  eggs. 

When  the  phylogeny  of  this  subfamily  is  studied,  there  becomes 
evident  the  primitive  character  retained  by  the  Lachnini.  In  these 
forms  the  beak  structure  and  the  nature  of  the  antennae  and 
cornicles  point  to  a  primitive  condition.  The  sexes,  too,  indicate 
this,  though  not  markedly  more  than  in  other  tribes.  But  the  fact 
that  these  forms  are  mostly  conifer  feeders  should  not  be  overlooked. 
It  is  the  opinion  of  the  writer  that  this  is  a  primitive  habit. 
The  Lachnus  branch,  therefore,  may  be  considered  the  lowest  branch 
of  the  Aphidinae.  If  the  wings  of  fossil  aphids  be  examined  it 
will  be  seen  that  by  far  the  greater  number  of  them  possess  a  wing 
structure  quite  different  from  that  of  our  living  forms.  The  radial 
sector  arises  back  of  the  stigma,  which  is  usually  very  long  and  narrow. 
This  character  is  retained  probably  only  in  the  Mindarinae.  It  is 
evident,  then,  that  during  the  development  of  the  present  Aphidinae 
this  vein  migrated  toward  the  tip  of  the  wing  until  it  came  to  stand 
either  in  the  middle  of  the  stigma  or  near  its  tip.  On  one  line  of 
this  migration  is  the  Lachnina  wherein  the  vein  has  reached  nearly 
to  the  tip  of  the  wing  and  become  short  and  straight.  The  remain- 
ing characters  apart  from  the  wings  have  in  these  forms  remained 
quite  primitive.  The  subtribe  Eulachnina  is  evidently  a  more 
specialized  group  on  this  same  line  of  development,  for  it  possesses 


GENERIC   CLASSIFICATION   OF   APHIDIDAE,  5 

the  same  type  of  wing.  Therefore  this  subtribe  is  considered  quite 
closely  related  to  the  subtribe  Lachnina  but  differing  from  it  in 
specialization  of  body  form,  cornicles,  and  eyes.  The  other  sub- 
tribes  of  the  Lachnini  have  quite  a  different  whig  structure.  While 
the  radial  sector  has  changed  its  position  considerably  from  that 
found  in  the  fossils  it  has  not  reached  the  tip  of  the  stigma  and  is  not 
straight,  but  much  curved.  In  this  regard,  therefore,  the  Pterochlor- 
ina  is  perhaps  the  most  primitive  subtribe  in  the  Lachnini,  although 
in  many  respects  it  is  specialized.  On  the  other  hand  there  are  two 
highly  specialized  subtribes,  the  Anoecina  and  the  Tramina.  It  is 
usually  the  custom  to  place  the  Anoecina  with  the  Pemphigina.  Its 
relations,  however,  are  here.  The  adult  forms  are  very  similar  in- 
deed to  the  lachnids.  The  sexual  forms,  on  the  other  hand,  are 
small  and  apterous  and  suggestive  of  the  sexes  of  the  Eriosomatinae, 
and  there  is  considerable  ground  for  placing  the  Anoecina  there. 
These  sexual  forms,  however,  seem  to  differ  quite  distinctly  from 
those  of  the  Eriosomatinae,  which  are  beakless  and  the  oviparous 
female  of  which  never  develops  more  than  one  egg.  The  develop- 
ment of  the  stigma  shows  quite  an  extreme  modification  from  the 
long,  narrow,  primitive  stigma. 

Near  this  same  line  of  development  is  the  rather  highly  specialized 
subtribe  Tramina.  The  most  marked  character  of  this  subtribe  is 
the  extreme  modification  of  the  hind  tarsi.  In  considering  only  the 
genus  Trama  it  might  be  thought  that  the  tribe  should  belong  with 
the  Eriosomatinae.  The  species  troglodytes  has  often  been  figured 
with  cornicles  in  the  apterous  form.  Specimens  from  Mordwilko 
and  Schouteden  determined  as  this  species  lack  them  and  the  writer 
therefore  considers  Trama  as  the  most  specialized  genus  in  the  sub- 
tribe.  Another  genus  represented  by  radicis  Kalt.  shows  cornicles 
very  large  and  of  a  typical  Lachnus  character.  Through  this  genus, 
therefore,  the  subtribe  can  be  placed  at  once  with  its  relatives  in  the 
Lachnini.  Apart  from  the  peculiar  tibial  character  this  genus  is  very 
lachnid-like. 

The  next  branch  from  the  Aphidinae  is  the  Callipterus  branch, 
which  may  be  considered  as  arising  somewhat  later  than  the  Lachnus 
branch.  From  this  offshoot  soon  after  it  arose  and  before  the  present 
genera  of  the  Callipterini  appeared  the  Thelaxini  separated. 

This  tribe,  the  old  Vacuini,  also  has  usually  been  placed  with  the 
Eriosomatinae.  There  are  some  resemblances,  it  is  true;  only  one 
egg,  for  instance,  is  usually  laid  by  the  sexual  female.  But  this  is 
not  always  the  case,  for,  according  to  Buckton,  more  than  one  egg  is 
sometimes  laid.  Such  a  condition  shows  that  the  one  egg  habit  is  of 
much  more  recent  development  than  in  the  Eriosomatinae.  More- 
over, the  sexual  female  is  very  different  in  structure.  She  is  not  the 


6  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

degenerate,  beakless,  norifeeding  individual  of  the  Eriosomatinae,  but 
feeds  upon  the  leaf  like  the  viviparous  forms.  Moreover,  in  external 
structure  these  insects  resemble  certain  ones  of  the  Callipterini,  to 
such  an  extent,  indeed,  that  Davidson  has  described  one  of  these 
forms  as  a  Chaitophorus.  Certain  of  the  structures  met  with  in  these 
forms  resemble  those  found  in  the  Hormaphidinae  and  on  these 
structures  the  tribe  might  be  placed  there.  The  writer  believes, 
however,  that  the  true  affinities  of  the  tribe  are  shown  by  comparison 
with  the  Phyllaphidina.  It  will  be  seen  at  a  glance  that  there  is  a 
very  close  resemblance  in  all  main  characters.  But  the  sexual  forms 
are  different,  although  not  so  strikingly  different  as  would  appear  at 
first.  In  the  Phyllaphidina  both  winged  and  wingless  ovipara  and 
as  a  rule  winged  males  occur.  But  in  some  species  (quercifoliae) 
intermediate  and  apterous  males  also  occur.  It  is  not  a  very  long 
call,  therefore,  from  the  apterous  males  and  females  of  the  Phyl- 
laphidina to  the  apterous  sexes  of  the  Thelaxini.  But  the  habit  of 
egg  laying  met  with  in  this  last  tribe  shows  that  it  has  been  on  this 
course  of  development  longer  than  has  the  Phyllaphidina.  The 
group  is  therefore  considered  as  a  tribe  which  has  separated  some- 
what earlier  and  yet  has  paralleled  in  some  ways  certain  characters 
of  the  Phyllaphidina. 

Continuing  with  the  Callipterus  branch  we  find  two  somewhat 
similar  lines  of  development,  the  one  represented  by  the  Callipterina 
and  the  other  represented  by  the  Chaitophorina.  Both  are  similar 
in  many  regards,  but  are  quite  different  in  the  armature,  particularly 
of  the  antennae. 

The  first  subtribe  separating  from  that  offshoot  represented  by 
the  Callipterina  is  the  Phyllaphidina.  This  seems  evident  from  the 
fact  that  the  oviparous  forms  of  some  species  are  yet  alate,  a  primi- 
tive condition  found  very  seldom  in  the  Aphididae.  The  next  off- 
shoot resulted  in  the  Callipterina  where  the  oviparous  forms  are  apter- 
ous, the  cornicles  of  moderate  development,  and  the  wing  veins 
usually  not  reduced.  From  this  offshoot  the  Saltusaphidina  evi- 
dently arose.  This  little  subtribe  is  closely  related  to  the  Callip- 
terina in  many  ways,  but  there  are  some  new  developments.  In 
the  first  place  the  power  of  leaping  has  become  developed  by  the 
enlarging  of  the  femora.  Secondly,  both  the  sexual  forms  have  lost 
their  wings,  which  the  male  usually  retains  in  the  Callipterina.  One 
of  the  most  important  points,  however,  is  the  fact  that  in  the  Sal- 
tusaphidina the  ocular  tubercles  which  represent  the  retained  larval 
eyes  are  absent,  whereas  they  are  quite  conspicuous  in  the  Callip- 
terina. 

On  this  same  Callipterus  branch,  but  somewhat  more  specialized 
than  the  Callipterina,  are  two  subtribes.  These  have  specialized 
in  opposite  directions,  the  one  toward  the  elimination  of  the  cornicles 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  7 

and  the  other  toward  the  development  of  them.  The  first  subtribe, 
the  Monaphidina,  lacks  cornicles  above.  The  second  of  these  two, 
the  Drepanaphidina,  possesses  them  in  varying  degrees.  In  this  last 
subtribe  the  males  are  winged,  and  the  females  have  developed  an 
extremely  long,  narrow  ovipositor. 

Coming  now  to  that  line  represented  by  Chaitophorus,  the  Chai- 
tophorina  are  found  to  be  the  most  generalized,  corresponding  quite 
closely  with  the  Callipterina.  In  this  subtribe  males  are  winged  as  a 
rule,  but  sometimes  in  the  same  species  they  are  intermediate  or 
apterous.  Arising  from  the  same  branch  with  the  Chaitophorina 
are  two  subtribes  specialized  in  different  directions,  like  the  sub- 
tribes  of  Callipterus.  Tlie  first,  Fullawayina,  lacks  cornicles  entirely, 
whereas  the  Pterocommina  has  developed  them  in  varying  degrees, 
as  has  the  Drepanaphidina.  This  concludes  the  subtribes  of  the 
Callipterus  branch. 

In  connection  with  these  insects  the  tribe  Greenideini  should  be 
considered.  The  cornicles  of  the  primitive  aphids  were  evidently 
small,  somewhat  rounded  or  conical,  and  armed  with  ban's.  In  the 
Greeliideini  the  insects  have  very  long  cylindric  or  somewhat  swollen 
cornicles  which  are  thickly  covered  with  prominent  hairs.  No  such 
well-developed  cornicles  are  met  with  in  any  of  the  other  tribes  of  the 
family,  although  they  are  approached  in  the  Macrosiphina.  In  this 
latter  subtribe  species  occasionally  occur  which  show  a  few  short  hairs 
on  the  cornicles.  It  seems  evident  then  that  the  Greenideini  separated 
from  the  Aphidinae  before  the  hairs  of  the  cornicles  disappeared. 
This  was  evidently  more  recent  than  the  development  of  the  tribe 
Lachmni  which  possesses  a  much  more  primitive  cornicle.  At  about 
the  same  time  that  the  ancestors  of  the  CaUipterini  separated  from 
the  Aphidinae,  other  forms  probably  separated  and  more  or  less 
paralleled  in  some  ways  the  ancestors  of  the  Macrosiphina,  but  unlike 
them  carried  the  hairs  of  the  cornicles.  They  thus  resulted  in  forms 
with  very  long  cornicles  similar  to  those  of  the  Macrosiphma  but 
armed  with  long  hairs.  In  other  characters,  too,  they  of  course  differ, 
particularly  in  regard  to  the  cauda. 

In  considering  the  further  development  of  the  Aphidinae,  a  more 
or  less  distinct  development  of  the  cornicles  and  antennal  tubercles  is 
found.  There  are  thus  two  types  which  separate  themselves,  rep- 
resented by  Aphis  and '  Macrosiphum  respectively.  These  may  be 
considered  as  leaving  the  aphid  line  at  about  the  same  time  after  the 
development  of  prominent  cornicles.  There  are,  consequently,  two 
subtribes,  the  Aphidina  and  the  Macrosiphina.  The  Cervaphidina 
represents  a  group  of  insects  armed  with  long,  somewhat  cylindric 
cornicles,  and  very  prominent  spinelike  protuberances.  The  number 
of  antenna!  segments  is  somewhat  reduced,  as  is  also  the  wing  sTena- 
tion.  It  seems  evident  then  that  this  is  a  subtribe  on  somewhat  the 


8  BULLETIN   826,   U.   S.  DEPARTMENT   OF   AGRICULTURE. 

same  line  of  development  as  the  Aphidina  but  developing  these  spec- 
ialized spines  during  the  same  period  in  which  the  wings  and  antennas 
have  become  reduced.  Still  another  subtribe,  the  Pentalonina, 
shows  a  very  peculiar  wing  venation.  This  is  not  so  much  a  primitive 
wing  as  a  more  specialized  one.  It  is  placed,  therefore,  as  one  of  the 
highest  subtribes  of  the  Aphidini. 

There  remains  yet  for  discussion  the  tribe  Setaphidini.  This,  it 
seems  evident,  belongs  with  the  Aphidinae.  In  regard  to  the  an- 
tennse  and  the  wings  it  is  quite  highly  specialized  but  in  regard  to  the 
cornicles,  cauda,  and  anal  plate  this  statement  can  not  be  made.  The 
natural  position  of  this  tribe  is  somewhat  doubtful.  Its  ancestors 
evidently  separated  from  the  aphidian  line  before  the  prominent  corn- 
icles of  the  Aphidina,  Macrosiphina,  etc.,  appeared  and  yet  the  species 
are  more  specialized  in  many  ways  than  are  members  of  those  sub- 
tribes.  It  would  appear  that  the  lines  separated  after  that  of  the 
Greenideini,  for  the  cornicles  are  not  hairy.  Yet  this  separation  must 
have  taken  place  a  considerable  time  before  that  of  the  Aphidina  and 
Macrosiphina.  The  tribe  is  placed,  therefore,  as  indicated  in  the 
diagram  (fig.  1). 

The  subfamilies,  other  than  the  Aphidinae,  include  the  most  spec- 
ialized members  of  the  family.  By  far  the  most  primitive  of  these 
subfamilies  is  the  Mindarinae.  This  subfamily,  as  has  been  indicated, 
is  a  remnant  from  the  past,  giving  some  idea  of  the  ancestors  of  the 
Eriosomatinae  and  the  Hormaphidinae.  The  wing  structure  is  partic- 
ularly worthy  of  study.  The  wing  of  no  other  living  aphid  is  like  it, 
but  this  peculiar  structure  is  abundantly  met  with  in  fossil  forms. 
The  media,  it  is  true,  is  more  reduced  than  in  certain  members  of  the 
Aphidinae,  but  this  is  of  very  little  importance  as  compared  with  the 
wing's  peculiar  structure.  The  form  also  feeds  upon  conifers  and  this 
is  undoubtedly  a  primitive  habit.  The  cauda  and  anal  plate  are 
unlike  those  met  with  either  in  the  Eriosomatinae  or  the  Hormaph- 
idinae. 

The  sexual  forms  are  interesting.  They  have  become  sufficiently 
specialized  toward  the  Eriosomatinae  to  have  lost  the  wings,  but  they 
retain  the  beak,  at  least  in  most  individuals,  and  feed.  The  ovaries 
of  the  oviparous  female  also  are  developed  so  that  a  number  of  eggs 
are  laid. 

The  two  remaining  subfamilies  are  the  most  highly  specialized  of  all 
aphids. 

The  Eriosomatinae  are  in  many  ways  more  specialized  than  the 
Hormaphidinae,  but  in  other  ways  they  are  more  primitive.  The 
whole  Eriosoma  line  separates  at  once  on  the  sexual  forms.  These 
are  small,  apterous,  and  beakless.  Throughout  their  life  they  take 
no  nourishment,  and  the  ovaries  of  the  oviparous  female  become 
atrophied,  so  that  only  one  develops  and  of  the  eggs  therein  only  one 


GENERIC    CLASSIFICATION    OF   APHIDIDAE.  9 

reaches  maturity.  The  most  primitive  tribe  on  this  line  is  the 
Eriosomatini.  The  forms  of  this  tribe  are  not  as  a  rule  distinct  gall 
formers.  They  possess  rather  prominent  cornicles  and  have  devel- 
oped special  wax  glands.  They  live  as  a  rule  upon  deciduous  trees, 
the  summer  forms  of  many  species  alternating  upon  the  roots  of 
plants. 

More  specialized  than  the  Eriosomatini  are  the  Pemphigini,  which, 
however,  are  very  similar  to  the  former  in  many  respects.  These  are 
distinct  and  true  gall  formers  on  deciduous  trees.  For  part  of  the 
year  they  are  usually  altogether  closed  within  the  gall.  Wax  secre- 
tion is  common  and  the  cornicles  are  present,  but  reduced  to  mere 
rings. 

The  Melaphini  are  closely  related  to  the  Pemphigini  and  are  gall 
formers  like  them.  These  forms,  however,  have  lost  entirely  the 
cornicles  which  are  usually  still  retained  in  the  Pemphigini.  \ 

A  somewhat  different  specialization  is  met  with  in  the  Prociphilini. 
Here  wax  secretion  has  developed  at  the  expense  of  the  cornicles  so 
that  these  organs  are  absent,  at  least  in  nearly  all  the  forms  of  the 
species.  Large  wax  plates  have  taken  their  places.  The  species  are 
not  true  gall  formers,  but  live  upon  foliage  which  they  cause  to  roll  or 
crumple  into  a  pseudogall.  Development  along  this  line  is  also  pres- 
ent in  the  next  tribe,  the  Fordini. 

Here  the  cornicles  are  also  absent,  being  replaced  by  large  wax 
glands,  but  the  species  are  nearly  all  root  feeders  and  are  usually 
associated  with  ants,  often  living  with  them  in  their  nests.  This  tribe 
may  be  considered  the  most  specialized  of  all  the  Eriosomatinae. 

The  same  specialization  in  the  sexual  forms  has  not  occured  in 
the  Hormaphidinae.  They  are  small  and  apterous,  it  is  true,  but 
they  possess  beaks,  they  feed,  and  the  oviparous  female  lays  more  than 
one  egg.  In  one  regard,  however,  these  insects  are  more  specialized. 
Many  of  them  have  developed  a  pecular  aleyrodiform  stage,  which  is 
quite  different  from  anything  occuring  elsewhere  in  the  family.  Along 
with  this  development  peculiar  wax  glands  have  made  their  appear- 
ance so  that  some  of  these  forms  look  very  much  like  aleyrodids 
and  are  indeed  often  mistaken  for  them. 

The  most  primitive  tribe  here  is  the  Oregmini,  which,  although  it 
possesses  many  of  the  other  characters  met  with  in  these  forms,  lacks 
the  aleyrodiform  stage.  These  insects  possess  quite  distinct  cornicles. 

Closely  related  to  the  Oregmini  is  the  Cerataphidini.  These  insects 
likewise  possess  cornicles  and  in  several  ways  suggest  the  Oregmini, 
but  they  have  developed  a  distinct  aleyrodiform  stage  and  in  this 
regard  are  much  more  advanced  than  the  members  of  that  tribe. 

Lastly,  and  perhaps  most  specialized  of  all,  are  the  Hormaphidini. 
These  insects  are  curious  gall  formers,  not  only  on  their  primary  host, 
but  often  on  their  secondary  one  as  well.  They  lack  cornicles  and 


10  BULLETIN   826,   IT.   S.   DEPARTMENT   OF   AGRICULTURE. 

have  developed  aleyrodiform  generations  and  wax  secreting  structures. 
In  many  ways  the  specialization  of  these  insects  is  most  remarkable. 

KEY  TO  THE  SUBFAMILIES  OF  THE  APHIDIDAE. 

1.  Sexual   forms   small  with   functioning  mouth   parts  absent.    Oviparous 

female  with  all  the  egg  tubes  present  or  indicated  in  the  embryo  but  the 
adult  possessing  only  one  tube  and  maturing  one  cell  so  that  one  egg  only 
is  laid.  Cornicles  much  reduced  or  absent.  Wax  glands  abundantly 
developed.  Wing  veins  usually  reduced.  Antennal  sensoria  prom- 
inent  ERIOSOMATIXAK. 

Sexual  forms  with  functioning  mouth  parts.     Nearly  all  the  ovarian  tubes 
developed  in  the  adult  oviparous  female .-. 2. 

2.  Radial  sector  of  forewing  inserted  mesad  of  the  stigma.     Sexes  small.     Ovi- 

parous female  laying  several  eggs MINDARINAE. 

Radial  sector  not  so  inserted  but  arising  from  the  stigma 3." 

3.  Forms  usually  gall  makers.     Wing  veins  much  reduced  so  that  the  media  is 

usually  simple.     Wax  glands  usual.     Antennal  sensoria  annular.     Aley- 
rodiform stages  common.     Sexes  wingless  as  a  rule  and  small.  .HORMAPHIDINAE. 
Forms  not  usually  gall  makers.     Wing  veins  often  not  reduced.     Wax 
glands  not  abundant.     Antennal  sensoria  oval  or  subcircular.     Aleyrodi- 
form stages  rare.     Cornicles  often  little  reduced .     Winged  males  common. 

APHIDINAE. 

Subfamily  I,  APHIDINAE. 

The  subfamily  Aphidinae  contains  many  of  the  most  primitive 
insects  in  the  family.  Indeed,  with  the  exception  of  the  Mindarinae 
the  subfamily  may  be  considered  as  by  far  the  most  primitive. 

The  oviparous  female,  in  all  the  tribes,  develops  the  ovaries  in  a 
normal  way  and  lays  several  eggs.  An  exception  to  this,  however,  is 
the  Thelaxini,  but  here  two  or  more  eggs  are  sometimes  laid.  The 
males  may  be  either  alate,  apterous,  or  intermediate,  and  in  many 
species  which  possess  the  migratory  instinct  they  are  often  produced 
on  quite  a  different  food  plant  from  the  oviparous  form.  The  stem 
mothers  are  in  practically  all  cases  apterous,  but  the  remaining  gener- 
ations throughout  the  year  may  or  may  not  be  winged.  In  many 
species  a  larger  percentage  of  winged  forms  occurs  in  certain 
generations  and  a  larger  percentage  of  apterous  forms  in  others. 
In  some  species,  however,  this  does  not  appear  to  be  the  case.  In 
certain  of  the  Callipterini  practically  all  of  the  viviparous  forms  other 
than  stem  mothers  are  winged. 

The  insects  are  mainly  foliage  feeders,  but  they  also  attack  the 
stems  and  roots.  They  occur  both  upon  woody  plants  and  herbs. 
Their  feeding  may  have  little  apparent  effect  upon  the  host  or  it  may 
cause  distortions  or  pseudogalls.  Some  species  are  particularly  in- 
jurious to  their  hosts  and  when  these  are  economic  plants  cause  much 
loss. 

Great  variation  is  met  with  amongst  the  members  of  the  subfamily. 
The  antennae  are  rather  long  and  slender  and  as  a  rule  are  armed 
with  subcircular  sensoria.  In  most  of  the  forms  the  sixth  segment 


GENEKIC   CLASSIFICATION   OF   APHIDIDAE.  11 

possesses  an  elongate  narrow  uuguis,  which  in  some  of  the  Callipterini 
and  Aphidini  is  remarkably  developed.  In  the  more  primitive 
groups,  however,  this  is  short  and  thumb-like.  The  head  of  the 
apterous  form  differs  much  from  that  seen  in  the  Eriosomatinae,  in 
that  true  compound  eyes  are  present  and  often  very  prominent,  and 
the  small  larval  eyes  are  seen  as  ocular  tubercles.  It  is  noteworthy 
that  in  the  Eriosomatinae  the  alate  forms  possess  distinct  compound 
eyes  but  the  apterous  forms  have  lost  them.  The  wings  are  in 
general  quite  similar  throughout  the  family  in  regard  to  the  venation. 
In  color,  shape,  arid  location  of  the  veins  there  is  often  considerable 
difference.  Moreover,  there  are  a  few  genera  amongst  the  different 
tribes  which  show  abnormal  wing  form,  of  which  genera  Microparsus 
is  a  good  example.  In  the  typical  forms  of  this  subfamily  the  media 
of  the  fore  wing  is  twice  branched,  but  it  is  very  commonly  branched 
only  once  and  it  is  rarely  simple. 

The  cornicles  show  remarkable  variation.  In  some  forms  of  the 
Callipterini  they  are  short  and  slightly  swollen  at  the  base,  in  the 
Lachnini  they  are  low  broad  cones,  whereas  in  the  Greenideini  they  are 
cylindrical  and  sometimes  longer  than  the  body.  Between  these 
extremes  every  gradation  occurs.  The  cornicles  may  be  straight  or 
they  may  be  swollen  to  a  greater  or  less  degree.  Practically  all  forms 
eject  a  colored  wax  from  these  organs  when  disturbed. 

The  cauda  shows  almost  as  much  variation  as  the  cornicles,  some- 
times being  short  and  rounded,  in  other  cases  elongate,  spatulate, 
or  conical,  and  in  others  distinctly  knobbed."  Variation  also  is  met 
with  in  the  anal  plate,  though  this  usually  is  rounded,  In  the  Callip- 
terini, however,  it  is  often  bilobed. 

The  tribes  of  the  subfamily  may  be  separated  as  follows: 

KEY  TO  THE  TRIBES  OF  THE  APHIDINAE. 

1.  Cornicles  situated  on  broad  flat  cones 2. 

Cornicles  truncate,  or  more  or  less  elongate 3. 

2.  Cornicles  and  antennae  hairy.     Antennae  with  the  unguis  short  and  thick .. 

LACHNINI. 

Cornicles  and  antennae   not  hairy.     Antennae  with  the  unguis  long  and 
slender SETAPHIDINI. 

3.  Cornicles  clothed  with  long  hairs GREENIDEINI. 

Cornicles  never  with  long  hairs 4. 

4.  Thorax  of  alate  form  with  the  lobes  not  prominently  developed;  oviparous 

form  small,  often  laying  one  egg.     Large  wax  plates  present THELAXINI. 

Thorax  of  alate   form  with  the   lobes  prominently  developed;  oviparous 
female  laying  several  eggs.     Large  wax  plates  usually  absent 5. 

5.  Cornicles  truncate  or  elongate;  when  elongate  the  cauda  knobbed,  and  the 

anal  plate  bilobed,  or  the  antennae'  prominently  hairy CALLIPTERINI. 

Cornicles  not  truncate,  usually  elongate.     Cauda  never  knobbed.     Anten- 
nae with  only  a  few  spinelike  hairs APHIDINI. 


12  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Tribe  LACHNINI. 

The  tribe  Lachnini  is  the  most  primitive  of  all  living  aphids,  with 
the  exception  of  the  Mindarinae.  The  genus  Mindarus  shows  in  its 
wing  structure  characters  more  primitive  than  any  of  the  Lachnini, 
but  in  other  characters  such  as  those  of  the  beak,  cornicles,  cauda, 
sensory  structures,  etc.,  the  Lachnini  are  very  primitive  insects.  In 
examining  the  fossil  wings  it  is  to  be  noted  that  the  radial  sector  is 
situated  back  of  the  stigma.  In  practically  all  living  aphids,  with  the 
exception  of  Mindarus,  this  vein  has  migrated  toward  the  tip  of  the 
wing.  In  primitive  forms  the  stigma  is  long  and  narrow,  whereas  in 
most  living  forms  it  has  become  more  or  less  compact.  In  the  subtribe 
Lachnina  the  radial  sector  has  become  a  very  short,  straight  vein  almost 
at  the  tip  of  the  wing.  This  shows  that  the  Lachnina  are  evidently 
more  advanced  than  the  Pterochlorina  in  which  the  radial  sector  is 
somewhat  curved  and  situated  near  the  middle  of  the  stigma.  The 
subtribe  Eulachnina  is  considerably  specialized,  as  indicated  by  the 
eyes,  the  shape  of  the  body,  and  the  cornicles.  It  is,  however,  as 
closely  related  to  the  Lachnina  as  are  any  of  the  other  tribes,  as  will 
be  seen  from  the  formation  of  the  wing.  The  Anoecina  hi  the  typical 
genus  shows  a  wing  with  a  short  blocky  stigma,  a  condition  quite 
different  from  that  seen  in  the  Lachnina,  and  the  radial  sector  is  here 
curved.  (In  Nippolachnus,  however,  the  stigma  is  still  long  and 
straight.)  Moreover,  the  sexual  forms  are  more  specialized,  being 
apterous  in  both  cases.  Anoecia,  therefore,  is  somewhat  removed 
from  Lachnus!"  The  genus  Trama  is  considerably  specialized,  in 
that  it  lacks  cornicles  in  the  apterous  form.  It  is,  however,  related 
to  Lachnus  through  Neotrama  with  small  cornicles,  and  Protrama 
with  large  hairy  cornicles. 

The  rostrum  in  the  Lachnini  is  in  many  species  five-segmented,  a 
primitive  character  most  marked  in  this  group.  The  freshly  emerged 
wing  of  a  lachnid  shows  that  M1?  M2,  and  M3+4  are  the  veins  repre- 
sented when  the  media  is  twice  branched,  and  that  in  some  species  no 
vein  is  formed  about  M2.  The  cubitus  and  first  and  second  anal 
are  present  in  the  forewing.  As  in  other  Aphididae,  however,  no 
vein  forms  about  the  second  anal.  The  radial  sector  is  in  Lachnus 
a  short,  straight  trachea  and  a  prominent  vein  forms  about  it.  The 
stigma,  as  in  all  members  of  the  family,  is  formed  by  radiusj.  In  the 
hindwing  both  media  and  cubitus  are  present  and  form  distinct 
veins. 

The  antennae  of  the  Lachnini  are  six-segmented  with  a  short 
unguis.  They  are  usually  armed  with  oval  or  subcircular  sensoria 
and  prominent  hairs.  In  fact,  the  entire  body  of  the  insect  is  hairy. 

The  cornicles  are  characteristic.  They  are  situated  on  distinct 
cones  which  are  constricted  before  the  somewhat  flanged  opening 
which  is  not  situated  over  the  center  of  the  cone.  The  cones  are  arm- 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  13 

ed  with  hairs.  Some  specialized  forms  have  small  cornicles  or  none 
at  all.  Wax-secreting  structures,  but  no  distinct  gland  areas,  are 
present  in  this  tribe  and  a  coating  of  fine  wax  is  often  found  over  the 
entire  insect,  including  the  appendages.  This  is  true  of  the  oviparous 
forms,  as  well  as  of  the  viviparous  ones. 

The  cauda  and  anal  plate  are  here  rounded,  never  developed  into 
elongate  structures  as  in  some  of  the  other  tribes  of  the  subfamily. 
The  sexual  forms  are  nearly  as  unspecialized  as  the  viviparous  ones. 
Both  sexes  possess  a  distinct  rostrum  and  take  food.  The  males  in 
the  typical  subtribes  are  winged.  The  females  are  apterous,  but  the 
ovaries  are  developed  and  several  eggs  are  laid  by  each  individual. 

.   KEY  TO  THE  SUBTRIBES  OF  THE  LACHNINI. 

1.  Radial  sector  of  fore  wings  curved  and  of  moderate  length 2. 

Radial  sector  of  fore  wings  short  and  straight,  situated  near  the  tip  of  the 

wing 4. 

2.  Hind  tarsi  extremely  elongate,  head  divided,  wing  venation  usually  faint. 

TRAMINA. 
Hind  tarsi  normal 3. 

3.  Stigma  short  and  thick,  sexes  both  apterous ANOECINA. 

Stigma  elongate,  males  often  winged PTEROCHLORINA. 

4.  Form  elongate  and  very  narrow;  antennae  with  bristles,  cornicles  not  hairy; 

eyes  without  ocular  tubercles EULACHNINA. 

Form  not  elongate ;  cornicles  on  hairy  cones ;  eyes  with  ocular  tubercles . .  LACHNIN  A  . 

Subtribe  ANOECINA. 

The  subtribe  Anoecina  is  suggestive  of  the  Tramina,  but  none  of 
the  forms  are  as  specialized  as  some  of  the  genera  of  that  subtribe. 
The  typical  genus  is  quite  distinctive  in  the  short  rounded  stigma 
and  in  the  sexual  forms.  The  genus  Nippolachnus,  however,  has 
a  stigma  quite  Lachnus-like  in  appearance.  Only  two  genera  are 
known  at  present. 

KEY  TO  THE  GENERA  OP  THE  ANOECINA. 

1.  Head  not  divided;  eyes  with  prominent  ocular  tubercles;  stigma  of  wing 

short  and  rounded Anoecia. 

2.  Head  divided;  eyes  without  ocular  tubercles;  stigma  long  and  straight. 

Nippolachnus. 

Genus  ANOECIA  Koch. 

Plate  1,  A-F,  I. 

1857.    A  noecia  Koch,  Die  Pflanzenlaiise  Aphiden,  p.  275. 

Characters. — Head  not  divided,  front  somewhat  rounded.  Eyes  prominent  but  not 
distinctly  set  off  from  the  head.  Antennae  of  six  segments,  armed  with  subcircular  or 
oval  or  elongate  sensoria  and  covered  with  hairs.  Fore  wings  with  the  media  once 
branched.  Stigma  short  and  thick.  Hind  wings  with  both  media  and  cubitus 
present.  Cornicles  situated  on  broad  hairy  cones.  Cauda  and  anal  plate  somewhat 
rounded. 

Spring  forms  free,  living  in  colonies;  summer  forms  often  subterranean.  Sexes 
small  and  apterous,  possessing  beaks  and  feeding.  Oviparous  female  laying  one  or 
more  than  one  egg. 

Type  (mono typical),  Aphis  corni  Fab. 


14  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Genus  NIPPOLACHNUS  Matsumura. 

Plate  I,  G,  H,  J,  K. 
1917.     Nlppolachnus  Matsumura,  Journ.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  382. 

This  genus  is  closely  related  to  Anoecia  Koch  and  yet  it  retains 
many  Lachnus  characters  which  are  not  present  in  Anoecia.  It  is 
separated  from  that  genus  by  several  important  points.  The  head 
is  distinctly  divided  as  it  is  not  in  Anoecia.  The  stigma  is  long  and 
Lachnus-like  and  the  eyes  have  not  the  small  prominent  ocular 
tubercles  of  Anoecia,  but  are  rounded  on  their  posterior  margins. 

Characters. — Antennae  of  six  segments,  armed  with  prominent  protruding  sensoria 
and  many  hairs;  head  divided;  eyes  without  prominent  ocular  tubercles.  Wings 
with  the  media  once  branched,  the  stigma  rather  long  and  narrow.  Cornicles  on  large 
broad  cones  entirely  covered  with  hairs. 

Spring  forms  free,  migrating  in  summer  to  alternate  hosts.     Sexes  small,  males 

winged. 

Type  (fixed  by  Matsumura,  1917),  Nippolachnus  piri  Mats. 

Subtribe  EULACHNINA. 

The  sub  tribe  Eulachnina  is  related  to  the  Lachnina  quite  closely, 
as  can  be  seen  by  the  wing  structure  where  the  radial  sector  is  a 
straight,  short  vein  extending  across  the  tip  of  the  wing.  The 
media  also  is  faintly  indicated.  In  the  other  branch  of  the  tribe  in 
which  the  Anoecina  and  Tramina  are  found,  the  radial  sector  is 
curved  as  it  is  in  Aphidina,  etc.  Members  of  the  Eulachnina  may, 
however,  be  separated  ai>  once  from  the  Lachnina  on  the  elongate, 
narrow  shape  of  the  body,  the  abruptly  rounded  cauda,  the  character 
of  the  cornicles,  and  the  absence  of  ocular  tubercles  upon  the  eyes. 
The  subtribe  is  evidently  quite  specialized  as  compared  to  the  Lach- 
nina. 

Characters. — Eyes  large  and  set  off  from  the  head;  ocular  tubercles  not  evident; 
antennae  slender,  armed  with  bristles  or  spines,  not  slender  hairs.  Cornicles  shallow, 
not  on  distinct  hairy  cones.  Cauda  abruptly  rounded.  Body  very  elongate  and 
slender,  scarcely  'wider  than  the  head. 

KEY  TO  THE  GENERA  OP  THE  EULACHNINA. 

1.  Antennae  of  five  segments,  armed  with  minute  bristles Essigella. 

Antennae  of  six  segments 2. 

2.  Media  once  branched,  antennas  with  long  stout  spines Eulachnus. 

Media  twice  branched Todolachnus. 

Genus  ESSIGELLA  Del  Guercio. 

Plate  I,  S-Y. 
1909.    Essigella  Del  Guercio,  Rivista  Patol.    Vegetale,  n.  s.,  v.  3,  p.  329. 

The  genus  Essigella  is  quite  similar  to  Eulachnus  with  the  excep- 
tion of  the  antennae. 

Characters. — Head  with  large  outstanding  eyes,  very  much  broader  than  long. 
Antennae  of  five  segments,  imbricated,  armed  only  with  a  few  minute  bristles.  Fore 
wings  with  the  media  faintly  indicated,  once  branched,  hind  wings  with  both  media 
and  cubitus  present.  Cornicles  chitinized  rings  situated  close  to  the  body,  no  hairy 
cones  present.  Cauda  rounded.  Body  elongate  and  narrow. 

Type  (mono typical),  Lachnus  calif arnieus  Essig 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  15 

Genus  EULACHNUS  Del  Guercio. 
Plate  I,  L-R. 

71853.   Cinaria  Curtis,  British  Entomology,  v.  12,  section  576. 
1909.  Eulachnus  Del  Guercio,  Rivista  Patol.  Vegetale,  n.  s.,  v.  3,  p.  329. 
1915.  Protolachnus  Theobald,  Bull.  Ent.  Res.,  v.  6,  p.  145. 

Del  Guercio  erected  the  genus  Eulachnus  without  setting  a  type, 
but  Wilson  1  has  indicated  agilis  Kalt.  as  the  type.  Apparently, 
therefore,  the  genus  must  be  based  upon  that  species.  Theobald's 
genus  was  based  on  his  tuberculostemmata,  a  species  in  which  the  char- 
acters are  the  same.  Cinaria  was  erected  with  pini  L.  as  type,  but 
this  was  questioned. 

Characters. — Head  divided,  eyes  rather  large  and  outstanding;  antennae  of  six  seg- 
ments, armed  with  long  stout  bristles.  Fore  wings  with  the  media  faintly  indicated 
and  once  branched ;  hind  wings  with  both  media  and  cubitus  present  but  faint;  cornicles 
minute  rings,  not  situated  on  hairy  cones.  Cauda  abruptly  rounded.  Body  elongate 
and  narrow. 

Type  (fixed  by  Wilson,  1911),  Lachnus  agilis  Kalt. 

Genus  TODOLACHNTJS  Matsumura. 
1917.     Todolachnus  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  381. 

The  writer  has  been  unable  to  study  the  type  of  this  genus,  but 
from  the  description  given  it  seems  to  represent  a  genus  belonging 
here  and  having  a  twice-branched  media.  The  description  of  the 
cornicles  as  "wart-like,  not  broader  at  base'7  would  indicate  its 
affinities  here,  also  the  words  "body  long,  nearly  parallel  on  the  lat- 
eral sides." 

Type  (fixed  by  Matsumura,  1917),  Todolachnus  abietis  Mats. 

Subtribe  LACHNINA. 

The  members  of  the  subtribe  Lachnina  may  be  separated  from 
those  of  other  subtribes  with  the,  exception  of  the  Eulachnina  by 
the  character  of  the  venation.  The  radial  sector  here  has  almost 
reached  the  tip  of  the  wing  and  become  a  short  straight  vein.  The 
genera  may  be  separated  as  follows: 

KEY  TO  THE  GENERA  op  THE  LACHNINA. 

1 .  Media  of  fore  wings  twice  branched Dilachnus. 

Media  not  twice  branched 2. 

2.  Media  once  branched 3. 

Media  simple Unilachnus. 

3.  Labium  lance-like Lachnus. 

Labium  obtuse Schizolachnus. 

Genus  LACHNUS  Burmeister. 

1835.    Lachnus  Burmeister,  Handbuch  der  Ent.,  v.  2,  pt.  1,  p.  91. 
1909.    Lachniella  Del  Guercio,  Redia,  v.  5,  p.  286. 

The  genus  Lachnus  Burmeister  was  erected  with  the  following 
included  species:  lapidarius  F&fo.,fagi  Linn.,  quercus  Linn.,  fasdatus 
Burm.,  and  punctatus  Burm. 

Of  these  species  the  following  were  removed  as  types  of  other 
genera:  fagi,  1857,  PTiyllaphis,  and  quercus,  1870,  Stomapkis. 

1  Wilson,  II.  F.  Xotes  on  the  synonymy  of  the  genera  included  in  the  tribe  Lachnini.    In  Ann.  Ent.  Soc. 
Amer.  v.  4,  p.  51-54,  1911. 


16  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

The  following  type  fixations  have  been  made  for  Lachnus: 

1840.  Aphis  roboris  Linnaeus,  Westwood. 

1863.  Lachnus  pinicola  Kaltenbacb,  Passerinl. 

1908.  A  phis  nudus  De  Geer,  Mordwilko. 

1910.  Lachnus  punctatus  Burmeister,  Wilson. 

1911.  Lachnus  fasciatus  Burmeister,  Wilson. 

Now  the  first  three  fixations  are  invalid,  since  the  species  were 
not  included  in  the  original  genus.  The  first  valid  fixation,  there- 
fore, is  that  of  Wilson,  1910,  when  he  set  punctatus  as  type.  This 
fixation,  according  to  present  rules,  can  not  be  changed  in  1911 
because  it  is  an  unfortunate  fixation,  but  punctatus  must  remain 
the  type  of  the  genus  Lachnus.  The  question  is  now  purely  zoo- 
logical. At  present  punctatus  is  unknown  and,  therefore,  the  genus 
Lachnus  must  remain  unknown  until  punctatus  is  discovered.  This 
is  the  situation,  if  the  rules  are  followed,  and  the  well-known  genus 
name  will  be  lost  to  us.  At  the  suggestion  of  numerous  aphid  workers 
we  are  holding  fasciatus  as  the  type  of  Lachnus  and  the  Commission 
will  be  asked  to  suspend  the  rules  in  this  case  on  account  of  the  long 
usage  of  the  name  Lachnus. 

In  1909  Del  Guercio  erected  the  genus  Lachniella  without  setting 
a  type  but  in  1911  Wilson  interpreted  this  genus  as  Lachnus  with 
fasciatus  Burm.  as  type.  Following  this  the  writer  definitely  desig- 
nated this  species  as  type.  Therefore,  Lachniella  will  become  a  syno- 
nym of  Lachnus. 

Characters. — Eyes  large,  with  distinct  ocular  tubercles  present.  Antennae  of  six 
segments  and  with  rather  prominent  hairs.  Cornicles  on  somewhat  shallow  hairy 
cones.  Fore  wings  with  the  radial  sector  short  and  straight ;  stigma  elongate ;  media  once 
branched.  Labium  lance-like. 

Type  (by  suspension  of  rules),  fasciatus  Burm. 

Genus  DILACHNUS  Baker. 
Plate  II,  A-C. 

1919.     Wilsonia  Baker,  Can.  Ent.,  v.  51,  p.  212. 
1919.    Dilachnus  Baker,  Can.  Ent.,  v.  51,  p.  253. 

Characters. — Eyes  with  distinct  ocular  tubercles.  Antennae  of  six  segments  and 
armed  with  slender  hairs  and  circular  sensoria.  Cornicles  on  rather  broad  hairy  cones. 
Fore  wings  with  radial  sector  straight,  media  twice  branched ;  hind  wings  with  both 
media  and  cubitus  present. 

Type  (fixed  by  Baker,  1919),  Lachniella  gradlis  Wlsn . 

Genus  SCHIZOLACHNUS  Mordwllko. 

Plate  II,  D. 
1908.    Schizolachnus  Mordwilko,  Ann.  Mus.  Zool.  1'Acad.  Imp.  des  Sci.,  St.  Petersbourg,  v.  13,  p.  375. 

The  genus  Schizolachnus  was  erected  by  Mordwilko  with  tomen- 
tosus  De  Geer  (pineti  Fab.)  as  type. 

Characters. — Eyes  large  and  with  ocular  tubercles  present.  Antennae  of  six  seg- 
ments and  with  rather  prominent  hairs.  Cornicles  on  somewhat  shallow  hairy  cones. 
Fore  wings  with  the  radial  sector  straight;  stigma  elongate;  media  once  branched. 
Labium  obtuse. 

Type  (fixed  by  Mordwilko,  1908),  Aphis  tomentosus  De  Geer. 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  17 

Genus  UNILACHNUS  Wilson. 

Plate  II,   E-G. 
1919.     Vnllachnus  V^'ilson,  Ent.  News,  v.  30,  p.  5. 

The  genus  Unilachnus  Wilson  appears  to  be  a  connecting  link 
between  this  subtribe  and  the  Eulachnina.  In  some  respects  the 
genus  is  very  suggestive  of  that  subtribe.  The  form  is  elongate  and 
the  cornicles  are  reduced.  They  are  armed,  however,  with  hairs  and 
are  not  so  reduced  as  in  the  Eulachnina.  The  ocular  tubercles  here 
are  very  rudimentary,  almost  absent,  and  in  this  regard,  too,  the  genus 
suggests  the  Eulachnina,  but  it  seems  to  belong  in  this  subtribe. 

Characters. — Form  elongate;  cornicles  somewhat  reduced;  ocular  tubercles  small; 
media  of  fore  wings  simple.  Antennae  of  six  segments  moderately  armed. 

Type,  Lachnus  parvus  Wilson. 

Subtribe  PTEROCHLORINA. 

The  genera  belonging  in  the  subtribe  Pterochlorina  are  in  some  ways 
more  primitive  than  those  of  the  Lachnina,  but  in  other  respects  some 
of  them  are  more  specialized.  The  radial  sector  of  the  fore  wings  is  still 
curved  and  in  some  genera  quite  elongate.  This  is  much  nearer  the 
early  type  of  wing  than  is  the  wing  of  the  Lachnina  where  the  radial 
sector  is  short  and  has  migrated  almost  to  the  tip  of  the  wing.  Of 
course,  the  distinct  curving  of  this  vein  found  in  some  of  the  genera 
is  an  advance  on  the  slightly  curved  elongate  vein  usually  met  with 
in  the  fossils,  but  to  our  mind  the  location  and  character  of  this  vein 
are  much  more  primitive  than  in  the  Lachnina.  The  stigmal  area  and 
the  sexual  forms  appear  considerably  more  primitive  than  in  the 
Anoecina,  the  specialization  of  which  has  been  in  a  different  direction 
from  that  of  the  Lachnina.  The  male  of  Stomaphis  is,  however,  an 
exception. 

Characters. — -Head  often  divided;  antennae  of  six  segments,  armed  with  hairs  and 
subcircular  sensoria.  Fore  wings  with  radial  sector  somewhat  curved  and  not  close 
to  the  tip  of  the  wing.  Cornicles  on  broad  hairy  cones.  Males  usually  winged. 

KEY  TO  THE  GENTERA  OF  THE  PTEROCHLORIXA. 

1.  Stigma  extending  along  costal  margin  almost  to  the  tip  of  the  wing..Longistigma. 
Stigma  not  so  extending 2. 

2.  Beak   extremely  long,  very   much   longer  than  body;  antennie  covered 

with  very  fine,  short  hairs Stomaphia. 

Beak  normal  in    length;  antennae  with  rather  stout  hairs,   often  quite 

long Pterochlorus. 

Genus  LONGISTIGMA  Wilson. 

Plate  II,  II-L. 

1909.    Longistigma  Wilson,  Can.  En< .,  v.  41,  p.  3S5. 
1909.    Davisia  Del  Gucrcio,  Redia,  v.  5,  p.  185. 

The  genus  Longistigma  Wilson  can  be  distinguished  at  once  by  the 
shape  of  the  stigma  which  is  drawn  out  at  the  tip  to  an  acute  point 
which  extends  almost  to  the  tip  of  the  wing.     The  type  species  is 
141613°— 120—  Bull.  826 2 


18  BULLETIN   826,   U.    S.   DEPARTMENT   OF   AGRICULTURE. 

very  large,  one  of  the  largest  aphids.  Del  Guercio  set  no  type 
for  his  subgenus  which  was  published  shortly  after  Wilson's. 
Wilson  placed  this  species  as  a  synonym  of  caryae  Harris  which  he 
made  the  type  of  Longistigma. 

Characters.— Size  large.  Head  somewhat  divided.  Eyes  large,  with  distinct 
ocular  tubercles.  Antennae  of  six  segments,  armed  with  subcircular  sensoria  and 
prominent  hairs.  Fore  wings  with  media  twice  branched,  radial  sector  not  a  great  way 
from  the  tip  of  wing  and  stigma  extending  around  almost  to  tip;  hind  wings  with  both 
media  and  cubitus present.  Cornicles  on  broad,  shallow,  hairy  cones.  Cauda  and  anal 
plate  somewhat  rounded. 

0  viparous  female  apterous.     Males  winged. 

Type  (monotypical),  Aphis  caryae  Harris. 

Genus  PTEROCHLORUS  Rondani. 

Plate  II,  S-X. 

1848.    Ptenchlorus  Rondani,  "Familia  Hemipterorum  Aphidinse'J  in  Nuovi  Annali  delle  Scienze 

Natural!,  p.  35. 
1855.    Dryobius  Koch,  Die  Pflanzenlause  Aphiden,  p.  225. 

1908.  Tuberolachnus  Mordwjlko,  Ann.  Mus.  Zool.  de  1'Acad.  Imp.  des  Sei.  St.  Petersbourg,  v.  13, 
p.  374. 

1909.  Dry  aphis  Del  Guereio,  Rcdia,  v.  5,  p.  262. 

1913.    Schizodryobius  Van  der  Goot,  Tidj.  voor  Ent.,  v.  56,  p.  130. 

1917.  Pterochlorid.es  Archangelsky,  Turkestan  Ent.  Stn.  Kept.  Tashkent. 

1918.  Tuberodryobius  Das,  Mem.  Ind.  Mus.,  v.  6,  p.  259. 

The  generic  name  Cinaria  was  used  for  Aphis  pini  L  ?,  and  Aphis 
roboris  L.  with  A.  pini  as  type.  In  the  writer's  opinion  Cinaria  can 
not  be  used  now  with  roboris  as  type.  Roboris  was  used  as  the  type 
of  Dryobius,,  therefore  this  name  is  clear.  Dryaphis  was  used  with 
Pterochlorus  as  a  synonym  but  the  name  as  a  generic  name  was  really 
first  used  by  Del  Guercio  in  1909.  Tuberolachnus  was  erected  in 
1908  with  viminalis  Boyer  as  type  but  the  difference  in  the  abdominal 
tubercle  is  not,  in  the  writer's  opinion,  sufficient  for  a  distinction. 
Van  der  Goot's  genus  is  plainly  a  synonym.  The  other  two  generic 
names  listed  were  used  with  LacTinus  persicae  Choi,  as  type.  This 
species  has  several  abdominal  dorsal  tubercles  but  for  the  same  reason 
as  Tuberolachnus  is  held  to  be  a  synonym.  . 

Characters.- — Head  somewhat  rounded.  Antennae  of  six  segments,  armed  with 
subcircular  sensoria  and  prominent  hairs.  Fore  wings  with  radial  sector  distinctly 
curved,  inserted  some  distance  from  tip;  media  twice  branched.  Hind  wings  with 
both  media  and  cubit  as  present.  Wings  often  banded  or  mottled,  cornicles  on  hairy 
cones.  Cauda  and  anal  plate  rounded.  Abdomen  sometimes  with  dorsal  tubercles. 
Type  (fixed  by  Rondani,  1848),  Aphis  roboris  Fab.  (=roboris  L.). 

Genus  STOMAPHIS  Walker. 
Plate  II,  M-R. 

1870.    Stomaphis  Walker,  The  Zoologist,  v.  28,  p.  2000. 
1881.    Rhynchoclcs  Altum,  Forst  Zool.,  v.  3,  p.  350. 

Of  the  five  species  originally  in  the  genus  Lachnus  the  species 
guercus  Linn,  was  removed  by  Walker  as  the  type  of  his  genus  Stoma- 
phis. Little  confusion  has  arisen  in  regard  to  this  species.  Altum's 
genus  was  erected  for  RJiynchocles  longirostris. 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  19 

Characters. — Head  slightly  rouuded;  eyes  large  with  distinct  ocular  tubercles. 
Antenme  of  six  segments  armed  with  large  subcircular  sensoria  and  thickly  covered 
with  fine  hairs.  Cornicles  situated  on  very  broad,  shallow,  hairy  cones.  Cauda  sub- 
conical,  slightly  rounded.  Anal  plate  rounded.  Fore  wings  with  radial  sector  rather 
long  and  somewhat  curved.  Media  twice  branched ;  hind  wings  with  both  media  and 
cubitus  present,  quite  widely  separated.  Beak  very  long,  much  longer  than  body. 
Males  wingless  and  with  rudimentary  mouth  parts. 

Type  (fixed  by  Walker,  1870),  Aphis  qitercus  Linn. 

Subtribe  TRAMINA. 

The  subtribe  Tramina  is  composed  of  insects  quite  specialized  in 
nature,  subterranean  and  often  associated  with  ants.  The  typical 
genus  Trama  is  the  most  specialized  of  all  and  is  in  some  char- 
acters suggestive  of  the  Fordina.  Its  relations  with  the  other 
Lachnini,  however,  are  shown  clearly  by  the  other  genera.  The 
genus  Trama,  as  described  by  Del  Guercio  (Redia,  v.  5)  possesses 
small  cornicles.  Specimens  of  troglodytes,  however,  received  from 
Schouteden,  Mordwilko,  and  others  lack  cornicles  entirely.  Some 
other  forms  possess  them  either  as  very  small  cones  or  as  large 
Lachnus-like  structures.  These  latter  are  evidently  the  most  primi- 
tive and  to  these  is  given  the  name  Protrama.  The  insects  with 
small  cornicles  are  grouped  under  the  name  Neotrama.  The  genera 
may  be  separated  as  follows : 

KEY  TO  THE  GENERA  OF  THE  TRAMINA. 

1.  Apterous  form  entirely  without  cornicles  and  with  rudimentary  eyes Trama. 

Apterous  form  with  cornicles 2. 

2.  Apterous  form  with  large,  broad,  Lachnus-like  cornicles  and  large  distinct 

compound  eyes Protrama. 

Apterous  form  with  small  cone-like  cornicles  armed  with  a  few  hairs  and 
more  or  less  rudimentary  eyes : Neotrama. 

Genus  PROTRAMA,  n.  gen. 
Plate  III,  P-T. 

Head  divided,  front  straight,  eyes  prominent  and  set  off  from  the  head.  Antenna? 
of  six  segments  armed  with  haira  and  small  subcircular  protruding  sensoria.  Cornicles 
situated  on  broad,  low,  hairy  cones.  Cauda  and  anal  plate  rounded.  Hind  tarsi 
extremely  elongate.  Wing  venation  faint;  fore  wings  with  the  media  twice  branched; 
hind  wings  with  both  media  and  cubitus  present. 

Type.   Trama  radicis  Kalt. 
Genus  TRAMA,  Heyden. 

Plate  III,  N. 

1837.     Trama  Hcydcn,  Mus.  Senkb.,  v.  2,  p.  293. 

Head  divided  but  not  prominently  so,  front  straight.  Apterous  form  with  the  eyes 
reduced  to  a  few  facets.  Antennae  of  six  segments;  cornicles  absent;  cauda  sub  conical, 
rounded.  Anal  plate  rounded.  Entire  insect  minutely  hairy. 

Type  (mono typical),   Trama  troglodytes  Heyden. 


20  BULLETIN   826,   U.   S.   DEPARTMENT  OF   AGRICULTURE. 

Genus  NEOTRAMA,  n.  gen. 
Plate  III,  M,  O. 

Head  somewhat  flat.  In  the  apterous  form  the  eyes  reduced ;  antennae  of  six  segments 
and  the  cornicles  on  very  small  cones  with  a  few  scattered  hairs.  Cauda  subconical, 
rounded.  Anal  plate  rounded.  Entire  body  covered  with  fine  hairs.  Hind  tarsi 
greatly  elongate. 

Type,  Trama  troglodytes  Del  Guercio  (=Neotrama  delguercioi  Baker). 

Tribe  THELAXINI. 

It  has  been  the  custom  of  most  writers  to  place  the  Thelaxini 
(Vacuini)  in  the  Eriosomatinae,  often  possibly  because  of  the  fact  that 
only  one  egg  is  laid  by  the  oviparous  female.  But  the  female  is  quite 
different  in  structure  from  the  beakless  females  of  the  Eriosomatinae 
and  the  other  forms  are  very  different  indeed. 

With  the  Hormaphidinae  there  are  more  resemblances,  the  most 
striking  of  which  is  the  structure  of  the  thorax.  The  mesothorax 
indicates  very  faintly  the  lobes  so  prominent  in  most  forms.  The 
presence  of  distinct  cornicles,  however,  is  very  different  from  the 
forms  in  the  Hormaphidinae  lacking  these  although  possessing  a  some- 
what similar  thorax. 

The  sensory  structures,  too,  are  widely  different,  being  similar  to 
those  found  in  the  Phyllaphidina.  Indeed,  the  antennae  are  very  like 
those  of  that  subtribe.  The  oviparous  forms  of  the  Phyllaphidina, 
however,  lay  several  eggs  and  may  be  either  winged  or  apterous  and 
the  males,  though  sometimes  apterous,  are  usually  winged.  The 
venation  of  the  Thelaxini  is  more  reduced  than  in  the  Phyllaphidina. 
Taking  all  of  these  facts  into  consideration  it  seems  evident  that  the 
Thelaxini  should  be  placed  in  the  Aphidinae  and  somewhat  related  to 
the  Phyllaphidina,  a  subtribe  which  belongs  in  the  Callipterini.  It  is 
evident,  however,  that  the  Thelaxini  must  stand  somewhat  apart; 
it  is  placed,  therefore,  as  a  tribe  of  the  subfamily  Aphidinae  next  to 
the  Callipterini.  In  this  tribe  the  specialization  of  the  ovipara  has 
advanced  beyond  that  of  the  Callipterini  in  that  only  one  egg  is  laid, 
but  according  to  Buckton  several  eggs  may  be  laid  and  the  distinct 
beak  is  evidence  of  relationship. 

Characters. — Cornicles  present  as  chitinized  rings  on  shallow  hairy  cones.  An- 
tennae somewhat  setose,  with  oval  or  subcircular  sensoria.  Cauda  somewhat  semi- 
circular or  distinctly  knobbed.  Body  usually  armed  Avith  hairs  or  etout  spines. 

Sexual  forms  small  and  apterous,  possessing  beaks;  oviparous  female  as  a  rule  laying 
only  one  egg. 

Forms  living  free  upon  the  foliage. 

The  genera  may  be  separated  by  the  following  key: 
KEY  TO  THE  GENERA  OF  THE  THELAXINI. 

Cauda  distinctly   knobbed Thelaxes. 

Cauda  not  knobbed  but  somewhat  semicircular Glyphina. 


GENERIC   CLASSIFICATION   OF  APHIDIDAE.  21 

Genus  GLYPHINA  Koch. 
Plate  III,  G-L 

1857.     Glyphina  Koch,  Die  Pflanzenlause  Aphiden,  p.  259. 
1911.     Travaresiella  Del  -Guercio,  Redia,  v.  7,  p.  299. 

Characters. — Corrncles  present  as  somewhat  elevated  rings.  Antenna  5-seg- 
mented,  minutely  setose,  armed  with  a  few  stout  hairs  and  somewhat  subcircular 
sensoria.  Fore  wings  with  the  media  once  branched;  hind  wings  with  only  the  media 
present.  Cauda  not  knobbed,  fomewhat  rounded,  anal  plate  rounded.  Body  covered 
with  hairs. 

Forms  living  upon  the  foliage  of  plants. 

Type  (monotypical),  Glyphina  betulae  Kalt. 

Genus  THELAXES  Westw. 
Plate  III,  A-F. 

Vacuna  of  authors,  not  Heydcn. 

1840.     Thelaxes  Westw.,  Int.  Mod.  Class.  Ins.  Synopsis,  v.  2,  p.  118. 

In  1837,  Heyden  erected  his  genus  Vacuna  based  on  coccinea 
Hey  den.  He  definitely  stated  that  he  thought  Phylloxera  Boyer  was 
the  same  genus.  Kaltenbach  stated  that  coccinea  is  a  Phylloxera 
and  so  considered  dryophila  as  type  of  Vacuna,  as  this  species  was  in- 
cluded in  the  genus  by  Heyden.  On  the  authority  of  Schouteden  and 
other  European  workers  coecinea  is  now  considered  a  Phylloxera  and 
another  type,  dryopJiilo,,  can  not  be  set  for  the  genus-  in  order  to  apply 
Vacuna  to  the  genus  as  now  understood.  Vaouna  with  coccinea  as 
type  will  become  a  synonym  of  Phylloxera  and  another  name  will 
be  necessary  to  apply  to  the  genus  having  dryophila  as  type.  The 
next  name  used  appears  to  be  Thelaxes  Westwood. 

Characters. — Cornicles  present  as  chitinized  rings  on  broad  lov/  cones.  Antennae 
of  the  stem  mother  5-segmented.  Alate  form  with  3-segmented  antennae,  sensoria  oval 
or  subcircular.  Fore  wings  with  the  media  once  branched,  hind  wings  with  the 
cubitus  lacking.  Cauda  distinctly  knobbed,  anal  plate  rounded.  Sexual  forms  small 
and  apterous,  possessing  distinct  beaks  and  feeding;  oviparous  female  producing 
normally  but  one  egg. 
Type  (fixed  by  Westwood,  1840),  Thelaxes  quercicola  Westw. (= Aphis  dryophila  Schr.) 

Tribe  CALLIPTERINI. 

The  tribe  Callipterini  is  composed  of  forms  which  live  upon  the 
foliage  of  plants.  The  species  in  many  of  the  sub  tribes  hare  developed 
peculiar  habits.  Some  forms  are  almost  solitary  whereas  others 
live  in  colonies.  Some  have  developed  the  power  of  leaping, 
while  others  are  sedentary.  The  sexual  forms  do  not  vary  greatly 
from  the  viviparous  forms.  In  nearly  all  of  the  subtribes  the  males  arc 
winged,  though  in  the  Saltusaphidina  they  are  apterous.  In  the  other 
tribes  intermediate  males  may  occur  in  the  same  species  with  alato 
males.  The  oviparous  females  are  nearly  always  apterous,  although  in 
the  Phyllaphidina  alate  ovipara  may  occur.  Both  sexes  feed  and  the 
ovaries  of  the  oviparous  female  are  developed  so  that  several  eggs  are, 
laid. 


22  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

The  wing  veins  are  not  reduoed  in  most  species.  In  some,  however 
the  radial  sector  of  the  fore  wings  is  very  faint  or  entirely  absent. 
This  condition  is  met  with  in  several  genera  of  the  Callipterina.  That 
it  is  this  vein  which  is  lacking  is  indicated  by  the  trachea  of  the  freshly 
emerged  wing.  Here  the  media  is  represented  by  M1?  M2,  and  M3+4. 
The  oubitus  and  first  anal  are  distinct  tracheae,  whereas  the  second 
anal  is  faintly  indicated.  In  the  hindwing  besides  the  radial  sector 
three  oblique  tracheae  are  present ;  these  are  the  media,  cubitus,  and 
first  anal.  Only  the  media  and  cubitus  are  represented  in  the  vena- 
tion. 

Considerable  variation  is  met  with  in  the  cornicles  of  this  tribe 
but  they  are  never  long  and  prominent  as  in  the  Aphidini.  The  usual 
form  is  the  truncate  one  represented  in  Myzocallis,  Chaitophorus,  etc. 
Very  often  the  cornicles  are  sculptured.  In  some  cases  they 
are  reduced  to  small  cup-shaped  structures  and  in  others  they  are 
represented  by  mere  rings. 

The  antennse,  as  a  rule,  are  long  and  slender  and  armed  with  few 
sensoria.  These  sensoria  are  usually  small,  subcircular  or  oval.  In 
rare  cases  they  are  somewhat  elongate. 

The  cauda  in  this  tribe  is  as  a  rule  knobbed  and  the  anal  plate 
bilobed.  In  some  cases,  ho\vever,  the  cauda  and  anal  plate  are  both 
rounded.  In  the  Saltusaphidina  the  anal  plate  is  divided  and  the 
cauda  remains  distinctly  knobbed. 

Wax  secretion  is  present  to  a  limited  extent  in  this  tribe.  It  is 
most  developed  in  the  Phyllaphidina.  Here  there  are  large  lateral  ab- 
dominal wax  plates  in  all  of  the  forms  and  the  insects  present  a  wool- 
like  appearance  on  the  foliage.  In  the  genus  Euceraphis  wax  secretion 
is  found  to  a  limited  extent.  In  one  species,  mucidus  Fitch,  it  is, 
however,  abundant  and  the  insects  of  this  species  often  seem  to 
float  in  the  air,  a  peculiar  appearance  cdmmon  also  in  the  Erioso- 
matinae.  In  the  Saltusaphidina  also  distinct  wax  plates  occur, 
particularly  in  the  oviparous  forms.  These  are  arranged  along  the 
abdominal  segments. 

The  habit  of  leaping  is  common  in  the  Saltusaphidina  as  the  name 
implies.  Here  the  muscles  of  the  femora  are  greatly  enlarged  for  this 
purpose.  Many  of  the  other  members  of  the  tribe  approach  this  con- 
dition, especially  in  the  genus  Monellia.  Others,  although  they  do  not 
distinctly  leap,  drop  so  suddenly  when  disturbed  that  they  almost  ap- 
pear to  leap  from  the  foliage.  Our  common  Symydobius  on  the 
birch  is  difficult  to  collect  on  account  of  such  a  habit  and  other  forms 
of  Callipterina  are  very  similar  in  action.  Certain  species  in  this 
tribe  are  closely  attended  by  ants  in  return  for  the  honeydew  excreted. 
Some  species  are  protected  by  these  Hymenoptera  by  means  of  sheds 
or  roofs  built  over  colonies  on  the  leaves  or  twigs.  These  sheds  are 
found  quite  commonly  upon  the  leaves  of  the  oaks  protecting  the  spe- 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  23 

cies  described  as  Symydobius  albasiphus  by  Davis  and  here  placed  as 
Neosymydobius. 

The  internal  structure  of  insects  of  this  tribe  appears  not  to  differ 
markedly  from  the  structure  in  other  groups.  Witlaczil,  however,  has 
reported  that  in  certain  members  of  this  tribe  the  intestine  forms  a 
closed  loop  almost  similar  to  that  found  in  the  Chermidae. 

As  a  rule,  in  this  tribe,  the  various  forms  met  with  in  the  subfamily 
occur.  In  the  genus  Monellia,  however,  in  some  species  at  least,  ap- 
•  terous  viviparous  forms  seldom  occur,  nearly  all  the  viviparous  forms 
being  alate. 

The  subtribes  may  be  separated  by  the  following  key: 

KEY  TO   THE  SUBTRIBES  OF  THE  CALLIPTERINI. 

1.  Eyes  with  ocular  tubercles  present,  head  not  elongate 2. 

Eyes  without  ocular  tubercles  present,  head  often  elongate SALTUSAPHIDINA. 

2.  Antennae  armed  with  rather  long,  prominent  hairs 3. 

Antennae  usually  only  with  minute,  sometimes  stout  bristles 5. 

3.  Cornicles  absent FULLAWAYINA. 

Cornicles  present 4. 

4.  Cornicles  cylindrical  or  vasiform , PTEROCOMMIXA. 

Cornicles  truncate,  enlarged  at  base CHAITOPHORINA. 

5.  Cornicles  absent  above MONAPHIDINA. 

Cornicles  present,  position  as  usual G. 

6.  Cornicles    reduced    to    mere    rings;    large    lateral  abdominal  wax   plates 

present PHYLLAPHIDIXA. 

Cornicles  usually  not  reduced  to  mere  rings;  no  large  abdominal  wax  plates 
present 7. 

7.  Cornicles   variable,  often   long  and   somewhat   swollen;   oviparous  female 

with  an  elongate  ovipositor DREPANAPHIDIXA. 

Cornicles  never  long;  always  short  and  truncate;  oviparous  fem?le  not  always 
with  an  elongate  ovipositor CALLIPTERIXA. 

Subtribe  PHYLLAPHDOINA. 

The  subtribe  Phyllaphidina  is  erected  for  the  species  related  to  the 
genus  Phyllaphis.  Many  of  the  characters  show  these  species  as 
quite  closely  related  to  the  Callipterina,  while  in  other  ways  they  very 
strongly  suggest  the  Thelaxini,  as  indicated  under  the  discussion  of 
the  tribe. 

Characters. — Cornicles  present;  antennae  of  six  segments,  minutely  setose,  sensoria 
elongate  or  subcircular;  cauda  knobbed  or  rounded,  anal  plate  often  bilobed,  wax 
glands  present.  Forms  living  free  or  in  pseudogalls.  Sexual  forms  often  alate,  some- 
times, however,  apterous  or  intermediate,  showing  that  the  apterous  condition  has 
developed  but  recently;  oviparous  female  producing  several  eggs. 

KEY  TO  THE  GENERA  OF  THE  PUYLLAPHIDINI. 

1.  Anal  plate  deeply  cleft  and  U-shaped Shivaphis. 

Anal  plate  entire  or  somewhat  bilobed,  not  deeply  cleft 2. 

2.  Cauda  rounded,  anal  plate  entire 3. 

Cauda  knobbed,  anal  plate  somewhat  bilobed Phyllaphis. 

3.  Oviparous  females  with  annular  sensoria Neophyllaphis. 

Oviparous  females  with  small  transverse  sensoria Taraalia. 


24  BULLETIN   826,   U.   S.   DEPARTMENT  OF   AGRICULTURE. 

Genus  NEOPHYLLAPHIS  Takahashi. 
1920.  Neophylla/Jiis  Takahashi,  Can.  Knt.,  v.  52,  p.  20. 

Cfutracters. — Cornicles  slightly  elevated.  Antennae  of  six  segments,  with  narrow, 
traasverse  sensoria.  Fore  wings  with  media  twice  branched,  hind  wings  with  both 
media  and  cubitus  present.  Cauda  and  anal  plate  rounded,  cauda  sometimes  slightly 
constricted.  Oviparous  females  winged  and  possessing  annular  sensoria.  Forms  wax 
secreting  and  living  free  on  the  plants. 

Type  (monotypical),  Neophyllaphis  podocarpi  Takahashi. 
Genus  PHYLLAPHIS  Koch. 

Tlate  IV,  FF,  GO. 
1857.    Phyllaphis  Koch,  Die  Pflanzenlause  Aphiden,  p.  248. 

The  well-known  genus  Phyllaphis  Koch  is  represented  by/agriL., 
but  as  indicated  under  Tamalia  has  been  made  to  include  species  of 
somewhat  different  structure. 

Characters. — Cornicles  present  as  chitinized  rings  which  are  very  slightly  elevated  on 
low  conical  bases.  Antennae  of  six  segments,  long  and  slender,  minutely  setose,  sen- 
soria narrowly  oval.  Fore  wings  with  the  media  twice  branched;  hind  wings  with  both 
media  and  cubitus  faintly  indicated.  Cauda  knobbed,  anal  plate  slightly  divided. 

Forms  living  upon  the  foliage,  sometimes  producing  a  curling  of  the  leaves.  Males 
usually  winged;  oviparous  form  apterous,  producing  several  eggs. 

Type  (monotypical),  Aphis  fagi  L. 

Genus  TAMALIA  n.  gen. 
Plate  IV,  HH,  II. 

The  genus  Tamalia  Baker  is  erected  for  Pemphigus  coweni  Ckll., 
a  species  which  has  since  been  placed  in  Phyllaphis.  It  is,  however, 
quite  distinct  from  the  type  of  the  genus  and  undoubtedly  represents 
a  new  genus.  Mr.  Theodore  Pergande  received  this  species,  coweni, 
from  California  and  made  some  notes  on  the  material,  thinking,  how- 
ever, that  it  was  a  new  species.  He  gave  it  a  provisional  name  and 
the  new  generic  name  here  used.  This  name,  as  far  as  our  knowledge 
goes,  never  was  published.  Among  the  large  number  of  new  genera 
conceived  by  Pergande  this  is  one  of  the  few  valid  ones  and  his  man- 
uscript name,  therefore,  is  used  here. 

Characters. — Cornicles  present  as  mere  flanges  on  low,  broad,  conical  bases.  Antennae 
of  six  segments  minutely  setose  and  with  narrow  sensoria.  Fore  wings  with  media 
once  branched,  hind  wings  with  both  media  and  cubitus  present.  Cauda  and  anal 
plate  both  rounded.  Abdominal  wax  plates  present. 

Forms  living  in  pseud ogalls.  Sexes  sometimes  both  winged,  oviparous  form  pro- 
ducing several  eggs. 

Type,  Pemphigus  cou-eni  Ckll. 

Genus  SHIVAPHIS  Das. 

1918.    Shivaphis  Das,  Mem.  Ind.  MILS.,  vol.  6,  p.  215. 

The  genus  Shivaphis  was  erected  with  celti  Das  as  type,  and  celti  is 
evidently  the  species  rcclescribed  as  ChromapJiis  celticolens  by  Essig.1 

1  ESSIG,  E.  O.,  and  KUWANA,  S.  I.  Some  Japanese  Aphididfe.  In  Proc.  Cal.  Acad.  Sci.,  v.  8,  no.  3, 
p.  95, 1918. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  25 

The  writer  has  studied  a  series'  of  specimens  of  this  species  taken  in 
1907  on  Celtis  sinensis.  There  seems  little  doubt  that  the  genus  is 
related  to  Phyllaphis.  The  dorsal  wax  glands  are  of  much  the  same 
structure  as  those  found  in  fagi  L.  The  deeply  cleft  anal  plate, 
however,  at  once  separates  the  two.  The  cauda  which  is  almost 
cylindrical  in  some  specimens  is  quite  distinctly  knobbed  in  others. 

Characters. — Cornicles  present  as  mere  rings.  Head  without  prominent  antennal 
tubercles.  Antennae  of  six  segments,  sensoria  elliptical.  Fore  wings  with  the  media 
twice  branched,  hind  wings  with  both  media  and  cubitus  present.  Cauda  cylindrical 
or  knobbed.  Anal  plate  deeply  divided.  Males  winged.  Oviparous  females  apter- 
oue.  Rows  of  wax  glands  present. 

Type  (monotypical),  Shivaphis  celli  Das. 

Subtribe  CALLIPTERINA. 

The  subtribe  Callipterina  is  a  somewhat  large  and  interesting 
one.  Some  of  the  species  are  very  large  and  more  or  less  solitary, 
others  are  small  and  live  in  colonies.  In  some  genera  apterous 
viviparous  forms,  with  the  exception  of  the  stem  mother,  seldom 
occur,  while  in  other  genera  they  are  as  a  rule,  present.  The  males 
are  in  most  cases  alate  and  the  oviparous  forms  apterous.  The  species 
of  nearly  all  the  genera  live  upon  the  leaves  of  plants.  They  do  not 
affect  these  greatly,  as  a  rule,  although  when  abundant  the  insects 
often  seriously  interfere  with  the  proper  development  of  the  trees 
attacked.  Many  of  the  insects  are  armed  upon  their  bodies  with 
prominent  spines  or  tubercles. 

Characters. — Cornicles  present,  truncate  in  form.  Antennae  with  setae  or  spines, 
of  six  segments,  and  armed  with  subciroular  or  in  a  few  cases  somewhat  elongate  sen- 
soria. Wings  often  clouded,  mottled,  or  banded.  Cauda  as  a  rule  knobbed,  anal 
plate  usually  more  or  less  indented  or  bilobed.  Body  often  armed  with  capitate 
spines  or  tubercles. 

KEY  TO  THE  GENERA  OF  THE  CALLIPTERINA. 

1.  Cauda  distinctly  knobbed,  anal  plate  usually  bilobed  or  sometimes  deeply 

divided 3. 

Cauda  not  distinctly  knobbed,  anal  plate  entire  or  almost  so 2. 

2.  Antennae  minutely  setose,  sensorium  at  base  of  unguis  oval  or  somewhat 

rounded ;  oviparous  female  with  secondary  sensoria Symydobius. 

Antennas  not  minutely  setose,  sensorium  at  base  of  unguis  long  and  narrow, 
oviparous  female  without  secondary  sensoria Euceraphis. 

3.  Anal  plate  deeply  divided  with  a  U-shaped  cleft  so  that  the  lobes  appear 

as  distinct;  cauda  knobbed Therioaphis. 

Anal  plate  bilobed,  not  deeply  divided ;  cauda  very  markedly  knobbed ...         4. 

4.  Antenna?  and  often  the  cornicles  with  prominent  hairs Callipterus. 

Cornicles  and  antennae  without  such  hairs 5. 

5.  Cornicles  much  reduced;  wings  sometimes  horizontal  in  repose Monellia. 

Cornicles  truncate,  fairly  well  developed;  wings  not  horizontal  in  repose 6. 


26  BULLETIN   826,   U.   S.   DEPARTMENT  OF   AGRICULTURE. 

6.  More  or  less  distinct  antennal  tubercles  present;  oviparous  female  with 

secondary  sensoria Calaphis. 

No  distinct  antennal  tubercles  present ;  oviparous  female  without  secondary 
sensoria 7. 

7.  Anal  plate  slightly  indented,  sometimes  almost  entire;  no  apterous  vivi- 

parous forms  developed Chromaphis. 

Anal  plate  distinctly  bilobed;  apterous  viviparous  forms  common Myzocallis. 

Genus  CALAPHIS  Walsh. 

Plate  IV,  S,  U. 

1863.     Calaphis  Walsh,  Proc.  Ent.  Soc.  Phila.,  v.  1,  p.  301. 

1913.    Siphonocallis  Del  Guercio,  Redia,  v.  9,  p.  293. 

1913.    Callipterindla  Van  der  Goot,  Tijdschr.  voor  Ent.,  v.  56,  p.  118. 

Walsh  erected  his  genus  Calaphis  for  his  betulella,  a  species  which 
lacks  the  radial  sector  in  the  wing,  and  on  this  character  he  based 
his  genus.  Del  Guercio  based  his  Siphonocallis  on  betulaecolens 
Fitch,  distinguishing  it  from  his  conception  of  the  genus  Callipterus, 
which  conception,  however,  was  not  according  to  type.  A  study  of 
betida-ecolens  shows  that  in  all  respects,  with  the  exception  of  the  radial 
sector,  this  species  is  similar  to  betulella.  Many  specimens  of  betula-e- 
colens  lack  the  radial  sector  and  in  most  cases  it  is  only  faintly  indi- 
cated at  best.  These  two  species,  therefore,  are  probably  congeneric. 

The  genus  Callipterinella  was  based  on  betularius  Kalt.,  and  this 
species  proves  to  be  very  similar  to  betulaecolens.  It  is  true  that  the 
frontal  tubercles  are  not  prominent  in  this  species  as  they  are  in  the 
type  of  Calaphis.  There  seems  no  doubt,  however,  that  all  of  these 
three  species  are  closely  related.  This  relation  is  shown  in  part  by 
the  sexual  forms.  The  oviparous  females  all  possess  sensoria  on  the 
antennas  and  are  very  similar  in  other  body  characters.  This  pres- 
ence of  sensoria  in  the  oviparous  form,  while  not  important  in  some 
groups,  separates  quite  distinctly  this  small  group  of  species  from  those 
of  the  Myzocallis  type.  It  seems  evident  that  the  relations  of  betu- 
larius are  with  betulaecolens  and  betulella.  Callipterinella,  therefore, 
is  also  a  synonym. 

Characters. — Cornicles  present,  distinct,  truncate.  Antennae  of  six  segments, 
armed  with  oval  sensoria  and  placed  on  more  or  less  distinct  tubercles.  Fore  wings 
with  the  media  twice  branched,  the  radial  sector  either  absent  or  faintly  indicated, 
sometimes,  however,  complete;  hind  wings  with  both  media  and  cubitus  present. 
Cauda  distinctly  knobbed,  anal  plate  bilobedj  body  with  prominent  hairs. 

Forms  living  more  or  less  solitary  upon  the  foliage,  sexes  notmarkedly  different  from 
the  other  forms;  oviparous  female  producing  several  eggs  and  possessing  sensoria  upon 

the  antennae. 

Type  (monotypical),  Calaphis  betulella  Walsh. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  27 

Genus  CALLIPTERUS  Koch. 
Plate  IV,  I,  J. 

1855.  Callipterus  Koch,  Die  Pflanzenlaiise  Aphiden,  p.  208. 

1870.  Callaphis  Walker,  The  Zoologist,  v.  5,  p.  2000. 

1881.  Ptychodes  Buckton,  Mon.  British  Aphids,  v.  3,  p.  39. 

1904.  Panaphis  Kirkaldy,  The  Entomologist,  v.  37,  p.  279. 

1917.  Nippocallis  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  365. 

In  erecting  the  genus  Callipterus,  Koch  included  a  number  of  species 
among  which  was  juglandis  Kalt,  In  I860,  Passerini  set  juglandis 
as  the  type  of  Callipterus  and  erected  his  Myzocallis  for  species  similar 
to  coryli  Goetz. 

Most  writers  overlooked  Passerini's  work  and  considered  the  genus 
Callipterus  in  the  light  of  Myzocallis.  This  is  the  conception  com- 
monly held  by  many  to-day.  The  application  of  Callipterus,  how- 
ever, must  be  restricted  to  species  essentially  like  juglandis.  All  the 
other  generic  names  listed  as  synonyms  were,  with  one  exception, 
used  with  this  same  species  juglandis  as  type  and  therefore  require 
little  comment. 

The  genus  Xippocallis  was  erected  with  Tcuricola  Mats,  as  type. 
Specimens  of  this  species  studied  by  the  writer  are  not  in  good  con- 
dition for  the  observation  of  the  anal  plate.  All  the  characters 
visible,  however,  indicate  that  this  species  is  a  Callipterus. 

Characters. — Cornicles  present,  truncate  in  form,  rather  prominent  and  often  armed 
with  long  hairs.  Antennae  of  six  segments,  armed  with  stout  hairs,  sensoria  usually 
oval.  Fore  wings  with  the  media  twice  branched,  hind  wings  with  both  cubitus  and 
media  present.  Veins  usually  bordered.  Radial  sector  often  faintly  indicated. 
Cauda  not  distinctly  knobbed  in  all  cases.  Anal  plate  bilobed;  body  usually  covered 
with  prominent  hairs. 

Forma  living  free  upon  the  foliage.  Sexual  forms  not  differing  markedly  from  the 
other  forms,  oviparous  female  producing  several  eggs. 

Type  (fixed  by  Passerini,  1860),  Aphis  juglandis  Frisch. 

Genus  CHROMAPHIS  Walker. 

Plate  IV,  O,  P. 
1870.    Chromaphis  Walker,  The  Zoologist,  v.  5,  p.  2001. 

The  genus  Chromaphis  was  erected  with  juglandicola  Kalt.  as  type. 
It  is  related  quite  closely  to  Monellia. 

Characters. — Cornicles  moderate  in  size,  somewhat  flanged.  Antennae  of  seven  seg- 
ments armed  with  oval  sensoria.  Fore  wings  with  the  media  twice  branched,  hind 
wings  with  media  and  cubitus  present,  wings  not  held  horizontally  in  repose.  Cauda 
knobbed,  anal  plate  slightly  indented.  Sexual  forms  somewhat  similar  to  the  vivip- 
arous ones.  Males  usually  winged.  Oviparous  female  with  the  ovaries  developed 
normally,  laying  numerous  eggs.1 

Forms  living  free  upon  the  foliage  usually  all  summer;  viviparous  generations 
winged. 

Type  (inonotypical),  Aphis  juglandicola  Kalt. 

i  Davidson  believes  as  many  as  30  may  be  produced  by  one  female. 


28  BULLETIN   826,   U.    S.    DEPARTMENT   OF   AGRICULTURE. 

Genus  THERIOAPHIS  Walker. 
Plate  IV,  K,  I,. 

1870.     Thcrioaphis  Walker,  The  Zoologist,  p.  1999. 

1905.  Kallistaphis  Kirkaldy,  Can.  Ent.,  v.  37,  p.  417. 

1906.  EucaTipterus  Schouteden,  Ann.  Ent.  Soe.  Belg.,  v.  .50,  p.  31.1 

1915.    Neocallipterus  Van  der  Goot,  Beitrage  zur  Kennt.  der  IIoll.  Blattlause,  p.  320. 

The  genus  Therioaphis  Walker  was  erected  with  ononidis  Kalt.  as 
type,  and  ononidis  has  been  shown  by  Theobald  to  be  the  common 
"yellow  clover  aphis,"  trifolii  of  Monell.  This  species  has  a  deeply 
cleft  anal  plate  quite  different  from  that  of  Myzocallis.  Eucallipterus 
was  erected  with  tiliae  L.  as  type,  &,  species  with  quite  similar  struc- 
ture. Eucallipterus,  therefore,  will  become  a  synonym.  Betulicola 
Kalt.  has  been  used  as  type  by  Kirkaldy  and  Van  der  Goot.  Accord- 
ing to  Das,  Van  der  Goot  considers  this  congeneric  with  trifolii. 
Therefore  Therioaphis  is  the  name  that  must  be  used. 

Characters. — Cornicles  truncate,  rather  constricted  mesad  of  apex.  Antennae  cf  six 
segments  without  prominent  hairs  and  armed  with  subcircular  sensoria.  Fore  wings 
with  media  twice  branched ;  hind  wings  with  both  media  and  cubitus  present.  Wings 
often  variously  marked.  Prothorax  rather  elongate;  cauda  knobbed*  Anal  plate 
deeply  bifid  so  that  two  long,  narrow  lobes  are  formed.  Body  often  with  prominent 
hairs. 

Type  (monotypical),  Aphis  ononidis  Kalt. 

Genus  EUCERAPHIS  Walker. 
Plate  IV,  Q,  Pv. 

1S70.    Euceraphis  Walker,  The  Zoologist,  p.  2001. 

1908.     CaUipteroides  Mordvrilko,  Ann.  Mus.  Zool.  1'Acad.  Imp.  des  Fci.  St.  Petersbourg,  v.  13,  p.  377. 

1913.    CaUipteroides  Van  der  Goo:,  Tijd.  voor.  Ent.,  v.  56,  p.  151. 

When  Walker  erected  his  genus  Euceraphis  with  betulae  L.  as  type 
he  had  in  mind  evidently  the  same  species  as  that  described  by  Koch 
under  the  same  specific  name,  and  thus  separated  species  of  this 
type.  Mordwilko  in  1900  erected  the  genus  CaUipteroides  with 
nigritarsus  Heyden  as  type.  Specimens  of  this  species  received  from 
Mordwilko.  show  that  the  species  he  had  was  the  betulae  of  Koch  or 
at  least  a  species  very  close  to  it.  This  would  then  make  CaUipte- 
roides a  synonym  of  Euceraphis.  In  1913  Van  der  Goot  used  the 
name  CaUipteroides  with  beiulae  Koch  as  type  and  his  placing,  there- 
fore, should  be  under  Euceraphis. 

Characters. — Cornicles  present,  truncate.  Antennae  of  six  segments,  long  and  slender, 
armed  with  rather  narrow  sensoria  usually  near  the  base  of  segment  III,  the  unguis 
of  segment  VI  usually  not  much  longer  than  the  base,  sensorium  at  the  base  of  unguis 
long,  oval,  and  fringed ;  more  or  less  distinct  frontal  tubercles  present.  Fore  wings  with 
the  media  twice  branched,  hind  wings  with  both  media  and  cubitus  present.  Cauda 
usually  knobbed  and  rather  large.  Anal  plate  usually  entire.  Abdomen  of  the  alate 
form  often  with  distinct  wax-producing  glands.  Forms  very  large  and  usually  solitary 
in  habit,  sexes  similar  to  the  other  forms;  oviparous  female  producing  several  eggs. 
Type  (fixed  by  Walker,  1870),  Aphis  betulae  (L.)  Walker  (=Callipterus  betulae  Koch). 

i  There  is  considerable  evidence  for  keeping  this  genus  distinct. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  2& 

Genus  MONELLIA   Oestlund. 

Plate  IV,  M,  N. 
1887.    AfoneUia  Oestlund,  Osol.  and  Nat.  Hist.  Survey  Minn.,  Bui.  no.  4,  p.  41. 

The  genus  Monellia  Oestlund  was  erected  for  caryeUus  Fitch  and 
only  one  species  was  included  in  the  genus  at  the  time.  Several  other 
species,  however,  were  made  synonyms  of  caryeUus  which  are  quite 
different  from  that  species  and  fall  into  other 'genera. 

Characters.  — Cornicles  present  as  mere  rings;  antennae  slender,  of  six  segments;  sen- 
soria  oval  or  subcircular;  head  broad  for  its  length;  prothorax  prominently  separated 
from  the  mesothorax.  Fore  wings  with  the  media  twice  branched ;  hind  wings  with 
both  media  and  cubitus  present.  Wings  often  held  flat  upon  the  back,  cauda  knobbed, 
anal  plate  bilobed.  Forms  living  solitary  upon  the  leaves,  sometimes  haying  the 
power  of  leaping.  Apterous  forms  rare.  Sexes  feeding;  oviparous  female  laying 
several  eggs. 

Type  (monotypical),  Aphis  caryella  Fitclu 

Genus  MYZOCALLIS  Pass. 
Plate  IV,  G,  H. 

1860.  Myzocallis  Passerinl,  Gli  Afldi,  p.  28. 

1860.  Pterocallis  Passerini,  Gli  Afidi,  p.  28. 

1894.  Subcallipterus  Mordwilko,  Varshava  Univ.  Izviestiia,  v.  8,  no.  58,  p.  53. 

1894.  Tuberculatus  Mordwilko,  Varshava  Univ.  Izviestiia,  v.  8,  no.  58,  p.  60. 

1913.  Callipterus  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  116. 

1915.  Tuberculoides  Van  der  Goot,  Beitrage  zur  Kennt.  der  Holl.  Blattlause,  p.  313. 

1917.  Acanihocallis  Matsumura,  Jour.  Coll.  Apr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  368. 

1917.  Takecallis  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  373. 

When  Passerini  erected  the  genus  Myzocallis  in  1860  he  placed 
coryli  Goetz  as  type.  Later  in  the  same  work  he  erected  his  genus 
Pterocallis  with  dlni  Pass,  as  type.  This  species  proves  to  be  very 
similar  indeed  to  coryli,  so  similar  in  most  of  the  characters  that  the 
writer  believes  the  genus  Pterocallis  to  be  a  synonym  of  Myzocallis. 
The  type  of  the  genus,  coryli  Goetz,  was  also  placed  as  the  type  of 
Callipterus  by  Van  der  Goot,  in  1913,  in  spite  of  the  fact  that  another 
type  for  that  genus  had  been  set  in  1860.  Callipterus  Van  der  Goot 
(1913)  is  therefore  a  synonym  of  Myzocallis.  The  species  alni  Pass, 
is  universally  considered  the  same  species  as  alni  Fab.,  and  this 
species  was  made  the  type  of  the  genus  Subcallipterus  by  Mordwilko 
in  1894.  Subcallipterus  Mordwilko,  1894,  is  therefore  a  synonym  of 
Myzocallis  for  the  same  reason  as  is  Pterocallis  Pass.,  1860.  The 
species  querceus  Kalt.  was  made  the  type  of  the  Tuberculatus  by 
Mordwilko  in  1894,  but  this  species  seems  too  closely  related  to  coryli. 
Tuberculatus,  therefore,  becomes  a  synonym.  The  species  quercus 
Kalt.  was  made  the  type  of  Tuberculoides  by  Van  der  Goot,  1915, 
and  this  is  quite  typically  a  Myzocallis.  A  number  of  genera  have 
been  erected  by  Matsumura  which  are  so  very  little  different  from  the 
type  species  that  they  are  listed  here  as  synonyms.  This  author 
follows  the  idea  of  proportions  of  the  antennal  segments  as  generic 
characters. 


30  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Characters. — Cornicles  truncate  without  a  very  distinct  neck;  antennae  of  six 
segments  armed  with  a  few  minute  bristles  and  oval  or  subcircular  sensoria.  Fore 
wings  with  the  media  twice  branched ;  hind  wings  with  both  media  and  cubitus  present. 
Cauda  knobbed;  anal  plate  bilobed,  not  divided;  body  usually  with  stout  hairs. 

Type  (fixed  by  Passerini,  1860),  Aphis  coryli  Goetz. 

Genus  SYMYDOBIUS  Mordwilko. 
Plate  IV,  DD,  EE. 

1894.  SymydMus  Mordwilko,  Varshava  Universitetskiia  Izviestiia,  v.  8,  no.  58,  p.  65. 
1917.    Yczocallis  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  p.t.  6,  p.  369. 
Symdobius  of  later  authors. 

The  genus  Symydobius  Mord.,  which  was  erected  for  obloTigus 
Heyden,  has  often  been  spelled  Symdobius  by  subsequent  writers. 
This  is  probably  due  to  the  erection  of  the  genus  in  a  Russian  publi- 
cation which  is  available  to  few  workers,  at  least  in  this  country. 
Specimens  of  this  species  studied  were  collected  by  Mordwilko  at 
Petrograd  and  Warsaw  and  by  Schouteden  at  Brussels.  The  species 
which  passes  under  the  name  of  oblongus  hi  America  is  quite  distinct, 
as  has  been  pointed  out  by  the  writer. 

Characters. — Cornicles  present,  truncate  or  with  an  evident  neck  and  on  a  broad 
low  base.  Antennae  of  six  segments  armed  with  numerous  delicate  hairs,  sensoria 
somewhat  oval  or  subcircular;  sensorium  at  the  base  of  the  unguis  not  long  and  nar- 
row, with  a  fringe  but  without  a  prominent  one,  cauda  semicircular,  anal  plate  similar 
in  shape,  sometimes  slightly  indented.  Fore  wings  with  the  media  twice  forked; 
hind  wings  with  both  media  and  cubitus  present,  somewhat  separated  at  the  base. 

Type  (monotypical),  Aphis  oblonga  Heyden. 

Subtribe  SALTUSAPHIDINA. 

The  subtribe  Saltusaphidina  is  separated  from  the  other  related 
ones  principally  on  the  nature  of  the  head.  The  most  important 
character,  possibly,  is  the  structure  of  the  eyes,  in  which  the  ocular 
tubercles  appear  to  be  wholly  lacking. 

Characters. — Forms  living  usually  in  damp  places  upon  the  foliage  of  sedges  and 
grasses,  narrow  elongate  bodies,  eyes  with  ocular  tubercles  lacking,  legs  often  modi- 
fied for  leaping.  Oviparous  forms  apteious,  somewhat  similar  to  the  viviparous 
forms,  producing  several  eggs. 

KEY  TO  THE  GENERA  OF  THE  SALTUSAPHIDINA. 

Head  considerably  elongate,  cornicles  cup  shaped,  legs  modified  for  leap- 
ing  Saltusaphis. 

Head  not  much  elongated,  cornicles  mere  rings,  legs  not  modified  for  leap- 
ing  Thripeaphis. 

Genus  THRIPSAPHIS  Gillette. 

Plate  IV,  X. 
1917.     Thripsaphis  Gillette,  Can.  Ent.,  v.  49,  p.  193. 

The  genus  Thripsaphis  was  separated  from  Saltusaphis  for  &o$i 
Gillette,  which  the  present  writer  had  included  in  that  genus,  and 
certain  other  similar  species. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  31 

Characters. — Cornicles  .present  as  slightly  elevated  rings;  antennae  of  six  segments 
armed  with  subcircular  sensoria.  Eyes  without  ocular  tubercles.  Fore  wings  with 
the  media  twice  branched,  hind  wings  with  the  cubitus  sometimes  absent,  cauda 
knobbed,  anal  plate  divided,  body  with  spinelike  hairs. 

Forms  living  free  upon  grasses  and  sedges  in  moist  localities.  Sexes  apterous, 
oviparous  female  producing  several  eggs. 

Type  (fixed  by  Gillette,  1917),  Brachycolus  balli  Gill. 

Genus  SALTUSAPHIS  Theobald. 

Plate  IV,  V,  W. 

1915.    Saltusaphis  Theobald,  Bull.  Ent.  Research,  v.  6,  pt.  2,  p.  138. 

Characters. — Cornicles  present,  cup-shaped  or  truncate;  antennae  of  five  segments, 
minutely  setose;  sensoria  small  and  subcircular.  Head  elongate,  ocular  tubercles 
absent.  Fore  wings  with  the  media  twice  branched;  hind  wings  with  the  cubitus 
usually  absent;  cauda  knobbed;  anal  plate  divided,  caudal  extremity  of  the  abdomen 
sometimes  bilobed;  body  covered  with  spines  which  are  often  modified  into 
different  shapes. 

Forms  living  more  or  less  solitary  upon  the  leaves  of  grasses  or  sedges  in  marshy 
regions.  Sexual  forms  apterous,  oviparous  female  laying  several  eggs. 

Type  (monotypical),  Saltusaphis  stir  pus  Theo. 

Subtribe  DREPANOSIPHINA. 

The  subtribe  Drepanosiphina  is  evidently  related  to  the  callip- 
terine  branch  of  the  tribe  rather  than  the  chaitophorine  one.  It 
has  specialized  in  the  opposite  direction  from  the  Monaphidina  in 
that  the  cornicles  are  more  or  less  prominently  developed.  It  would 
appear  to  bear  the  same  relation  to  this  branch  of  the  tribe  as  does  the 
Pterocommina  to  the  chaitophorine  one.  There  is  considerable 
variation  in  the  development  of  the  cornicles,  even  within  certain 
of  the  genera.  Some  have  very  large  cornicles,  others  have  small 
ones.  They  all  appear,  however,  to  have  the  same  general  structure. 

Characters. — Cornicles  present,  varying  greatly  in  development  from  very  small  to 
very  large.  Cauda  somewhat  knobbed,  anal  plate  slightly  indented.  Oviparous 
female  with  a  long  drawn  out  ovipositor. 

KEY  TO  THE  GENERA  OF  THE  DEEPANOSIPHINA. 

1.  Cornicles  extremely  long  and  somewhat  swollen  in  the  middle Drepanosiphum. 

Cornicles  not  extremely  long  and  larger  atthe  base 2. 

2.  Cornicles  very  small,  truncate Neosymydobius. 

Cornicles  large  with  a  swollen  region  at  the  base Drepanaphis. 

Genus  DREPANAPHIS  Del  Guercio. 
Plate  IV,  JJ-LL. 

1909.    Drepanaphis  Del  Guercio,  Ri vista  di  Patologia  Vegetale,  n.  s.,  v.  4,  no.  4,  p.  49-50. 
1909.    Phymatosiphum  Davis,  Annals  Ent.  Soc.  America,  v.  2,  p.  196. 

The  two  generic  names  given  were  both  used  with  the  same  type 
species,  and  therefore  no  discussion  in  regard  to  the  use  of  the  names 
is  necessary. 

Characters. — Cornicles  large  but  not  of  the  same  shape  as  those  of  Drepanosiphum, 
being  rather  narrow  toward  the  distal  extremity  and  swollen  at  the  base.  Antennae 
of  six  segments  armed  with  subcircular  or  oval  sensoria  and  a  few  scattered  hairs. 
Fore  wings  with  the  media  twice  branched,  hind  wings  with  both  media  and  cubitus 
present.  Cauda  knobbed,  anal  plate  somewhat  indented.  Forms  living  more  or  less 


32  BULLETIN   826,   U.   S.    DEPARTMENT   OF   AGRICULTURE. 

solitary  upon  the  foliage  of  trees.     Males  winged.     Oviparous  female  with  a  distinct 
elongated  ovipositor  and  producing  several  eggs. 

Type  (monotypical),  Drepanosiphum  acerifolii  Thos. 

Genus  DREPANOSIPHUM  Koch. 
Plate  IV,  MM. 

1855.    Drepanosiphum  Koch,  Die  Fflanzenlaiise  Aphidcn,  p.  201. 
1885.    Type  fixation,  Lichtenstein,  Monographic  des  Aphidiens,  p.  175. 

Cornicles  very  long,  quite  distinctly  swollen  in  the  middle  or  subcylindric.  An- 
tennae of  six  segments  with  short,  scattered  hairs  and  oval  or  subcircular  sensoria. 
Fore  wings  with  the  media  twice  branched.  Hind  wings  with  both  media  and  cubitus 
present.  Cauda  knobbed,  anal  plate  slightly  indented. 

Forms  living  upon  the  foliage  of  plants,  males  usually  winged.  Oviparous  female 
with  a  distinct  elongated  ovipositor. 

Type  ffixed  by  Lichtenstein,  1885),  Aphis  platanoides  Schr. 

Genus  NEOSYMYDOBIUS,  n.  gen. 

The  genus  Neosymydobius  is  erected  for  species  similar  to  that 
described  as  Symydobius  albasipTius  Davis.  It  is  evident  that  this 
species  is  not  a  Symydobius.  Members  of  that  genus  are  very  large 
and  differ  in  several  ways.  We  place  the  present  genus  here  with 
considerable  doubt. 

Characters. — Cornicles  small,  truncate,  Callipterus-like.  Antennae  of  six  segments 
which  are  armed  with  a  few  rather  stout  hairs.  Fore  wings  with  the  media  twice 
branched,  hind  wings  with  both  media  and  cubitus  present.  Cauda  knobbed,  anal 
plate  slightly  indented.  Forms  living  in  colonies  upon  the  foliage  of  trees.  Body, 
particularly  of  the  apterous  forms,  covered  with  rather  stout  spine-like  hairs.  Ovipar- 
ous female  with  a  long  ovipositor  and  depositing  several  eggs.  Males  usually  winged. 

Type,  Symydobius  albasiphus  Davis. 

Sub  tribe  MONAPHIDINA. 

The  sub  tribe  Monaphidina,  erected  for  the  genus  Monaphis,  is  very 
similar  in  most  respects  to  the  Callipterina,  but  lacks  the  cornicle 
dorsally.  It  seems  evident  that  it  is  a  specialization  from  insects  of 
the  Callipterus  type  in  much  the  same  way  that  Fullawaya  is  related 
to  those  of  the  Chaitophorus  type.  One  genus  only  is  known  at 
present.  This  differs  more  from  the  Callipterina  than  does  Fullawaya 
from  the  Chaitophorina. 

Genus  MONAPHIS  Walker. 

1870.    Monaphis  Walker,  The  Zoologist,  p.  2001. 

1894.    Bradyaphis  Mordwilko,  Varshava  Univcrsitetskiia  Izviestiia,  v.  8,  p.  59. 

Few  remarks  on  the  synonymy  of  the  genus  Monaphis  Walker  are 
necessary  as  the  same  species  was  used  as  type  in  both  cases  men- 
tioned. The  genus  is  a  very  remarkable  one,  being  peculiar  in  many 
ways. 

Characters. — Cornicles  faint;  antennje  of  six  segments,  without  distinct  hairs, 
eensoria  small  and  circular;  cauda  somewhat  rounded  but  with  an  acute  point  or  pro- 
jection; anal  plate  similar;  fore  wings  with  the  media  twice  branched ;  hind  wings  with 
both  media  and  cubitus  present. 

Type  (fixed  by  Walker,  1870),  Aphis  antenna  in  Kalt. 


GENERIC    CLASSIFICATION    OF   APHIDIDAE.  33 

Subtribe  CHAITOPHORINA. 

The  sub  tribe  Chaitophorina  is  composed  of  aphids  which' are  similar 
in  many  ways  to  the  Callipterina.  They  differ  in  that  they  are  always 
armed  with  long  hairs  which  quite  prominently  cover  the  antenna  as 
well  as  the  other  parts  of  the  body.  Some  of  the  species  perhaps  are 
given  more  to  living  in  colonies  than  are  the  Callipterina,  but  this 
habit  varies  in  that  subtribe  as  well. 

KEY  TO  THE  GENERA  OF  THE  CHAITOPHORIXA. 

1 .  Cauda  quite  distinctly  knobbed 2. 

Cauda  not  knobbed  but  rounded 3. 

2.  Antennae  of  five  segments Sipha. 

Antennae  of  six  segments Chaitophorus. 

3.  Body  elongate ;  small  dimorphic  forms  developed Periphyllus. 

Body  not  elongate ;  no  dimorphic  forms  developed 4. 

4.  Antennae  of  five  segments Atheroides. 

Antennae  of  six  segments 5 

5.  Anal  plate  entire  or  slightly  indented Xeothomasia. 

Anal  plate  divided  into  two  quite  separate  parts Patchia. 

Genus  ATHEROIDES  Haliday. 

1839.  Atheroides  Haliday,  Ann.  and  Mag.  Xat.  Hist.,  v.  2,  p.  189. 

More  careful  collecting  of  marsh-inhabiting  species  may  show  that 
this  genus  is  a  specialization  from  the  Chaitophorina  as  is  the  Saltu- 
saphidina  from  the  Callipterina,  for  most  specimens  falling  here  seem 
to  lack  ocular  tubercles. 

Characters. — Antennae  of  five  segments  armed  with  stout  spines.  Fore  wings  with 
media  twice  branched;  hind  wings  with  both  media  and  cubitus  present.  Cornicles 
reduced  to  mere  rings.  Cauda  broadly  rounded.  Form  elongate  and  flat.  Entire 
insect  prominently  spined.  Species  living  on  sedges  and  grasses. 

Type  (set  by  Kirkaldy,  1906),  Atheroides  stmdatus  Ilaliday. 

Genus  CHAITOPHORUS  Koch. 

Plate  IV,  CC. 

1854.  Chaitophorus  Koch,  Die  Pflanzenlause  Aphiden,  p.  1. 

1870.  Tranaphis  Walker,  The  Zoologist,  p.  1999. 

1870.  A  rctaphis  Walker,  The  Zoologist,  p.  2000. 

1912.  Eichochaitophorus  Esslg,  Pom.  Coll.  Journ.  Ent.,  v.  4,  p.  721. 

1912.  Micrella  Essig,  Pom.  Coll.  Journ.  Ent.,  v.  4,  p.  716. 

1856.  Type  fixation,  Gerstaecker,  Bericht  for  1854,  p.  162. 

Ill  1854  Koch  erected  the  genus  Chaitophorus  with  several  species. 
Aphis  populi  was  made  the  type  by  Gerstaecker  in  1856.  The  same 
species  was  used  by  Walker  as  type  of  his  genus  Arctaphis  and  there- 
fore this  name  will  become  a  synonym.  The  types  of  both  of  Essig's 
genera  show  that  these  vary  little  from  populi.  The  genus  Tranaphis 
was  erected  with  salicivorus  Walker  as  type  and  this  species  is  similar 
in  general  characters  to  -populi.  Therefore,  Tranaphis  will  become 
a  synonym. 

141013°— -20—  Bull.  820 3 


34  BULLETIN   $26,   u.   S.   DEPARTMENT  OF   AGRICULTURE. 

In  1860  Passerini  used  cuceris  L.  as  the  type  of  Chaitophorus  and 
this  placing  has  often  been  followed,  but  that  of  Gerstaecker  has 
priority. 

Characters. — Cornicles  present,  truncate,  rather  prominent.  Antennae  of  &ix  seg- 
ments, armed  with  subcircular  sensoria  and  rather  prominent  hairs.  Fore  wings  with 
the  media  normally  twice  branched,  hind  wings  with  both  media  and  cubitus  piesent. 
Cauda  distinctly  knobbed.  Anal  plate  entire,  sometimes  somewhat  indented.  Sex- 
ual forms  not  differing  markedly  from  the  vi viparous  ones.  Males  winged,  as  a  rule, 
but  sometimes  intermediate  or  apterous.  Oviparous  females  apterous  with  the  ovaries 
normally  developed  and  producing  several  eggs.  Both  sexes  feeding. 

Forms  living  usually  upon  the  leaves  of  trees;  no  dimorphic  forms  developed. 

Type  (fixed  by  Gerstaecker,  1856),  Aphis  populi  L. 

Genus  PAT  CHI  A,  n.  gen. 

Characters. — Cornicles  truncate;  antennae  of  six  segments,  hairy  and  with  circular 
sensoria.  Fore  wings  with  the  media  twice  branched,  the  radial  sector  absent  or 
faintly  indicated;  hind  wings  with  both  media  and  cubitus  present.  Cauda  rounded 
or  slightly  conical;  anal  plate  divided  into  two  separate  parts. 

Type,  Patfhia  rirginiana  Baker. 

Patchia  virginiana,  n.  sp. 

Alate  viviparous  female. — Antennae  as  follows:  III,  0.48  mm.,  with  an  even  row  of 
about  12  subcircular  sensoria;  IV,  0.288  mm.;  V,  0.24  mm.;  VI  (0.16-0.192  mm.). 
Color  brown  with  a  large  black  patch  on  dorsum  of  abdomen  and  with  lateral  patches 
of  same  color.  Wings  with  the  radial  sector  absent  and  the  veins  heavily  bordered. 
Apterous  form  almost  solid  velvety  black.  Both  forms  secreting  wax. 

Found  on  the  bark  of  chestnut  at  East  Falls  Church,  Va.  The  type  is  in  the  U.  S. 
National  Museum  (Cat.  No.  23063). 

Genus  PERIPHYLLUS  Van  der  Hoeven. 
Plate  IV,  A  A,  BB. 

1852.  Phittophorus  Thornton,  Proc.  Ent.  Soc.  London,  n.  s.,  v.  2,  p.  78. 

1858.  Chelymorpha  Clark,  The  Microscope. 

1863.  Periphyllus  Van  dor  Hoeven,  Tijd.  voor.  Ent.,  v.  6,  p.  7. 

1913.  Chaitophorinella  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  150. 

1917.  Arakawana  Matsumura,  Journ.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  375. 

"In  1852  Thornton  used  the  name  Phillophorus  with  his  testudinatus 
as  type.  This  name  had,  however,  been  used  in  1840.  Koch  erected 
the  genus  Chaitophorus  in  1854  and  included  therein  a  number  of 
species.  In  1856  Gerstaecker  set  Aphis  populi  L.  as  the  type  of 
Chaitophorus  and  therefore  prevented  the  use  of  the  name  for  spe- 
cies such  as  testudinatus  unless  all  of  Koch's  species  are  included. 
In  1858  Clark  used  the  name  Chelymorpha  with  the  specific  name 
'pliyllopliora.  The  species  he  discussed  is  the  testudinatus  of  Thorn- . 
ton.  The  generic  name  Chelymorpha,  however,  was  used  as  early 
as  1834  and,  therefore,  is  not  available.  In  1863  Van  der  Hoeven 
employed  the  generic  term  Periphyllus  with  his  species  testudo  as 
type.  This  name  is  a  synonym  of  testudinatus  Thornton,  and  the 
generic  name  seems  to  be  the  first  one  available. 

In  1913  Van  der  Goot  employed  the  generic  name  Chaitophorinella 
with  testudinatus  as  type,  and  this  name,  therefore,  will  become  a 
synonym  of  Periphyllus. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  35 

Characters. — Cornicles  present,  truncate  in  form,  often  sculptured.  Antennae  of 
six  segments  (with  the  exception  of  the  dimorph)  armed  with  oval  sensoria  and  promi- 
nent hairs.  Fore  wings  with  the  media  twice  branched;  hind  wings  with  both 
media  and  cubitus  present.  Cauda  and  anal' plate  rounded. 

Forms  living  upon  the  foliage  of  trees.  Sexes  not  strikingly  different  from  the 
other  forms,  possessing  beaks  and  feeding.  Males  winged,  oviparous  females  with 
the  ovaries  normally  developed,  thus  laying  several  eggs.  Small  lamellate  or  hairy 
dimorphic  forms  produced  in  summer. 

Type  (monotypical),  Periphyllus  testudo  Van  der  Hoeven  (=testudinatus  Thorn- 
ton). 

Genus  NEOTHOMASIA,  n.  n. 

Plate  IV,  Y,  Z. 
1910.     Thomasia  Wilson,  Can.  Ent.,  v.  42,  p.  386. 

Wilson  erected  the  genus  Thomasia  with  populicola  Thos.  as 
type,  and  his  description  appeared  in  December,  1910.  The  same 
name  had,  however,  been  used  for  a  genus  of  Diptera,  the  description 
of  which  appeared  in  September,  1910.  A  new  name,  Neothomasia, 
therefore,  is  necessary  for  Wilson's  genus. 

Characters. — Cornicles  present;  antennae  of  six  segments  armed  "with  subcircular 
sensoria  and  prominent  hairs.  Fore  wings  with  the  media  twice  branched,  hind  wings 
with  both  media  and  cubitus  present.  Cauda  and  anal  plate  both  rounded. 

Forms  living  in  colonies  upon  the  leaves  or  bark  of  trees;  no  dimorphic  forms  pro- 
duced; sexual  forms  not  markedly  different  from  the  viviparous  ones.  Oviparous 
females  laying  several  eggs. 

Type  (monotypical),  Chaitophorus  populicola  Thos. 

Genus  SIPHA  Pass. 

1860.    SipTia  Passerini,  Gil  Afldl,  p.  29. 

This  genus  and  Atheroides  are  distinct  from  the  other  genera  in 
the  sub  tribe  by  possessing  five-segmented  antennae  instead  of  six- 
segmented  ones.  The  genus  has  not  been  much  confused  excepting 
by  Thomas's  placing  of  rubifolii.  For  a  time  some  workers  in  this 
country  were  led  to  conceive  of  the  genus  as  indicated  by  that  spe- 
cies which  in  reality  belongs  in  the  Aphidini. 

Characters. — Cornicles  present,  truncate,  short,  almost  mere  rings.  Antennae  of 
five  segments  armed  with  large  circular  seneoria.  Body  form  flat,  entire  insect  cov- 
ered with  rather  long  stout  hairs.  Fore  wings  with  the  media  twice  bran  died,  hind 
wings  with  both  media  and  cubitus  present.  Cauda  knobbed,  anal  plate  rounded. 
Forms  living  upon  the  leaves  of  grasses  usually  in  moist  localities,  sometimes  even 
submerged,  the  water  appearing  to  affect  them  little. 

Type  (fixed  by  Passerini.  1SGO),  Aphis  glyccriae  Kalt. 

Subtribe  PTEROCOMMINA. 

The  subtribe  Pterocommina  is  composed,  of  bark-feeding  insects, 
some  of  which  retain  quite  primitive  characters.  It  is  the  writer's 
opinion,  however,  that  they  are,  as  a  group,  more  specialized  than 
the  Chaitophorina,  but  closely  related.  This  is  indicated  by  the 
development  of  the  cornicles  met  with  in  the  species.  Like  the 


36  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Drepanosiphina,  this  development  varies  to  a  great  extent  in  the 
different  species. 

Only  two  genera  occur  in  the  tribe.     They  may  be  separated  as 

follows : 

KEY  TO  THE  GENERA  OF  THE  PTEROCOMMINA. 

Cornicles  cylindrical Pterocomma. 

Cornicles  somewhat  swollen Melanoxantherium. 

Genus  PTEROCOMMA  Buckton. 
Plate  IV,  P P. 

1857.  Cladobius  Koch,  Die  Pflanzenlause  Aphidcn,  p.  251. 

1860.  ApUoides  Passerini,  Gli  Audi,  p.  28. 

1879.  Pterocomma  Buckton,  Monog.  Br.  Aphides,  v.  2,  p.  142. 

1905.  Aristaphis  Kirkaldy,  Can.  Ent.,  v.  37,  p.  416. 

In  1857  Koch  erected  the  genus  Cladobius  with  populeus  Kalt.  as 
type.  This  name,  however,  had  been  used  previously.  So  Passerini 
in  1860  employed  the  name  Aphioides.  This  name  had  also  been 
used.  Kirkaldy,  therefore,  gave  the  new  name  Aristaphis  in  1905. 
In  1879,  however,  Buckton  described  the  genus  Pterocomma  with  a 
very  similar  species  as  type. 

Characters. — Cornicles-  present,  rather  short  and  cylindrical.  Antennae  of  six 
segments  armed  with  prominent  hairs  and  sub  circular  sensoria.  Fore  wings  with  the 
media  twice  branched;  hind  winga  with  both  media  and  cubitus  present.  Cauda 
and  anal  plate  rounded. 

Type  (monotypical),  Pterocomma  pilosa  Buckt. 

Genus  MELANOXANTHERIUM  Schouteden. 
Plate  IV,  NN,  OO. 

1879.     Jfelanoxanthus  Buckton,  Monog.  Br.  Aphides,  v.  2,  p.  21. 

1901.     Melanoxantherium  Schouteden,  Ann.  Ent.  Soc.  Belg.,  v.  45,  p.  113. 

In  1879  Buckton  described  the  genus  Melanoxanthus  with  solids 
L.  as  type,  but,  as  this  name  was  preoccupied,  Schouteden  suggested 
the  name  Melanoxantherium. 

Characters. — Cornicles  present,  variable  in  size,  but  usually  more  or  less  swollen. 
Antennae  of  six  segments  armed  thickly  with  hairs  and  possessing  oval  or  sub  circular 
sensoria.  Fore  wings  with  the  media  twice  branched,  hind  wings  with  both  media 
and  cubitus  present.  Cauda  and  anal  plate  rounded. 

Forms  living  in  colonies  usually  on  the  bark  of  trees,  males  usually  winged.  Ovi- 
parous females  laying  several  eggs. 

Type  (monotypical),  Aphis  salitis  L.1 

Subtribe  FULLAWAYINA. 

The  sub  tribe  Fullawayina  is  related  somewhat  closely  to  the 
Chaitophorina  from  which  it  is  a  specialization,  as  is  evidenced  by  the 
reduction  and  total  lack  of  the  cornicles.  It  is  very  suggestive  of 
Monaphis,  but  evidently  arose  from  quite  a  different  line  of  develop- 
ment, following  Chaitophorus  rather  than  the  Callipterus  group. 
Only  one  genus  is  represented. 

1  The  writer  is  forced  to  change  his  view  that  Pterocomma  pilosa  is  closely  related  to  populifoliae 
Fitch.  This  was  based  on  Pergande's  published  statement  of  his  examination  of  the  type.  Later  notes 
on  the  type  indicated  that  it  resembles  populea. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  37 

Genus  FULLAWAYA  Essig. 
1912.    Fullawaya  Essig,  Pomona  C!oll.  Journ.  Ent.,  v.  4,  p.  716. 

The  genus  Fullawaya  seems  to  bear  somewhat  the  same  relation  to 
Chaitophorus  that  Monaphis  does  to  Myzocallis.  Were  it  not  for  the 
hairy  condition  of  the  antennae  and  the  character  of  the  cauda  and 
anal  plate,  Fullawaya  might  he  placed  as  a  synonym  of  Monaphis, 
hut  it  is  evidently  unrelated. 

Characters. — Corniclea  absent;  antennae  of  six  segments,  armed  with  rather  long 
bristle-like  hairs,  sensoria  small  and  circular,  cauda  rounded  and  armed  with  long 
curved  hairs.  Fore  wings  with  the  media  twice  branched,  hind  winga  with  both 
media  and  cubitus  present. 

Forms  living  upon  the  roots  of  plants. 

Type  (monotypical),  Fullawaya  saliciradicis  Essig. 

Tribe  GREENIDEINI. 

The  tribe  Greenideini  was  first  separated  by  Wilson  under  the 
name  Trichosiphina.  The  insects  falling  here  show  a  most  remarkable 
development  of  the  cornicles.  These  are  sometimes  as  long  as  the 
entire  body  in  the  alate  forms.  In  the  apterous  individuals  they  are 
usually  swollen.  In  both  they  are  very  thickly  covered  with  long 
hairs,  a  condition  not  met  with  in  any  of  the  other  Aphidinae  with 
long  cornicles.  It  is  true  that  some  species  of  Macrosiphum  and  occa- 
sionally species  of  the  other  genera  show  here  and  there  minute  hairs 
on  the  cornicles,  but  they  do  not  approach  in  any  way  members  of 
this  tribe  in  cornicle  armature.  The  development  of  the  cauda  in 
this  group  is  also  remarkable. 

Characters. — Cornicles  present  and  remarkably  developed  into  cylindrical  or  slightly 
swollen  tubes  often  as  long  as  the  body  and  thickly  covered  with  long  hairs.  Antennae 
of  five  or  six  segments  armed  with  oval  or  subcircular  sensoria.  Males  winged.  Forms 
living  free  upon  the  foliage. 

KEY  TO  GEXESA  OF  GREKXIDEIXI. 

1 .  Antennae  of  five  segments Eutrichosiphum. 

Antennae  of  six  segments 2. 

2.  Fore  wings    with    the  media    twice    branched,    hind  wings    with  both 

media  and  cubitus  present Greenidea. 

Fore  winga  with  media  once  branched,  hind  wings  with  neither  media 
nor  cubitus  present Greenideoida. 

Genus  GREENIDEA  Schouteden. 

riato  V,  F-K. 

1905.  Greenidea  Schouteden,  Spol.  Zcylan,  v.  2,  p.  181. 

1906.  Trichosiphum  1'ergandc,  Ent.  News,  v.  17,  p.  206. 

In  1905,  Schouteden  erected  a  genus  for  the  SipJionopTiora  artocarpi 
of  Westwood  and  redescribed  the  species,  giving  details  lacking  in 
Westwood's  paper.  Pergande  erected  his  genus  Trichosiphum, 
making  his  anonae  the  type.  The  characters  Pergande  used  to  sep- 
arate his  genus  were  in  reality  those  of  Greenidea. 


38  BULLETIN  826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Characters. — Cornicles  extremely  long  and  hairy;  antenna?  of  six  segments  armed 
\vith  oval  or  subcircular  senspria  and  distinct  hairs.  Fore  wings  with  media  twice 
branched;  hind  wings  with  both  media  and  cubit  us  present.  Sexes  winged. 

Type  (monotypical),  Kiphonophora  artocarpi  Westw. 

Genus  GREENLDEOIDA  Van  der  Goot. 

Plate  V,  L-P. 

1900.     Greenidea  Wilson,  not  Schoutcden,  Ann.  Ent.  Soe.  Amer.,  v.  3,  p.  317. 
1916.    Greenideoida  Van  der  Goot,  Zur  Kenntniss  der  Blattliiuse  Java's,  p.  140. 

In  discussing  the  genera  of  the  Trichosiphini  in  1910,  Wilson  based 
his  descriptions  and  key  on  specimens  in  the  collection  of  the  Bureau 
of  Entomology.  Material  hi  that  collection  determined  as  artocarpi 
by  Pergande  proves  not  to  be  that  species  for  it  does  not  agree  with 
the  descriptions  given  either  by  Westwood  or  Schouteden.  'Wilson, 
therefore,  used  the  two  generic  terms  Trichosiphum  and  Greenidea. 
In  reality  the  species  listed  as  artocarpi  was  undescribed  at  that  time 
and  the  species  of  Trichosiphum  presented  all  of  the  characters  of 
Greenidea.  Since  that  time  Van  der  Goot  has  erected  the  genus 
Greenideoida  for  such  species  as  that  understood  by  Wilson  to  be 
artocarpi. 

Characters. — Cornicles  present,  very  long,  subcylindrical,  and  armed  with  long 
hairs.  Antennse  of  six  segments  armed  with  oval  or  eubcircular  sensoria.  Fore  wings 
with  the  media  once  branched ;  hind  wings  reduced  in  size  and  lacking  both  the 
media  and  cubitue.  Cauda  and  anal  plate  rounded. 

Type  (fixed  by  Van  der  Goot,  1916),  Gremideaida  elongata  V.  d.  <ioot. 

Genus  EUTRICHOSIPHUM  Essig  &  Kuwana. 

Plate  V,  A-E. 
1918.     EutrldiosipJium  Essig  &  Kuwana,  Proc.  Cal.  Acad.  Sci.,  v.  8,  no.  3,  p.  97. 

The  genus  Eutrichosiphum  Essig  and  Kuwana  was  erected  for  the 
species  pasaniae  Okj . 

Characters. — Similar  in  general  characters  to  Greenidea.  Antenna;  of  five  segments) 
armed  with  long  hairs  and  somewhat  oval  sensoria.  Cornicles  very  long,  subcylindrical 
and  covered  with  hairs.  Fore  wings  with  the  media  twice  branched,  hind  wings  with 
both  media  and  cubitus  present.  Cauda  and  anal  plate  rounded. 

Type  (monotypical),  Tricho#iphum  pasaniae  Okj. 

Tribe  SETAPHIDINI. 

The  correct  position  of  the  tribe  Setaphidini  is  somewhat  difficult 
to  ascertain.  In  some  ways  it  closely  resembles  the  Aphidini,  and 
in  others  suggests  the  Lachnini. 

It  has  Aphis-like  antennae  with,  however,  a  reduced  number  of 
segments.  The  venation  of  the  wings  is  also  reduced.  On  the  other 
hand  it  possesses  cornicles  situated  on  low  flat  cones  somewhat  like 
those  of  the  Anoecina  or  Lachnina.  But  these  cones  are  devoid  of 
hairs. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  39 

It  would  appear  that  this  tribe  separated  from  the  aphid  line  after 
the  cornicles  had  lost  their  armature  and  before  their  development 
as  indicated  in  the  Aphidina  and  Macrosiphina  began.  These  organs 
then  remained  somewhat  primitive  whereas  reduction  took  place  in 
the  antennae  and  wings.  One  genus  only  is  represented. 

Genus  SETAPHIS  Van  der  Goot. 

Plate  V,  Q-X. 

1916.    Setaphis  Van  der  Goot,  Zur  Kenntniss  der  Blattlause  Java's,  p.  153. 

Characters. — Cornicles  present  as  rings,  situated  on  low  broad  cones.  Antennae  of 
five  segments  armed  with  small  circular  sensoria.  Fore  wings  with  the  media  once 
branched,  hind  winga  reduced.  Cauda  and  anal  plate  rounded.  Body  with  two 
prominent  caudal  fingerlike  projections. 

Type  (fixed  by  Van  der  Goot,  1916),  Setaphis  luteus  V.  d.  Goot. 

Tribe  APHIDINI. 

The  tribe  Aphidini  is  by  far  the  largest  tribe  of  the  living  Aphididae. 
Many  of  the  most  common  species  in  the  family  as  well  as  many 
of  the  most  injurious  ones  belong  here,  and  it  is  these  forms  which 
correspond  to  the  popular  conception  of  the  family. 

Besides  being  abundant  they  are  varied,  and  a  large  number  of 
genera  is  therefore  found  in  this  tribe.  Specialization  has  taken 
place  in  a  number  of  directions,  but  particularly  in  the  development 
of  cornicles,  cauda,  etc.  The  wings  have  become  somewhat  reduced 
in  certain  genera,  but  as  a  rule  little  reduction  in  these  organs  has 
occurred,  the  venation  in  most  cases  being  as  complete  as  in  even 
the  most  primitive  forms  of  the  family.  The  antennae  have  developed 
an  elongate  filamentous  process  to  the  distal  segment,  which  in  the 
Lachnini  is  represented  by  a  very  short  thumblike  projection.  Wax 
secretion  is  found  scarcely  at  all  apart  from  that  produced  and 
secreted  by  the  cornicles.  The  head  shows  certain  peculiar  develop- 
ments in  some  of  the  tribes  in  that  the  antennae  are  situated  on 
prominent  tubercles  variously  shaped  and  armed. 

As  a  rule  the  body  is  more  or  less  naked,  being  covered  only  by  a 
few  scattered  hairs.  In  the  peculiar  specialized  Cervaphidina, 
however,  large  toothed  processes  extend  outward  from  the  body 
surface. 

Migration  between  a  primary  host  and  one  or  more  secondary 
hosts  often  occurs.  Apterous  and  alate  viviparous  forms,  therefore, 
are  common,  but  no  definite  relation  exists  between  them.  The 
forms  feed  mostly  upon  the  leaves  of  trees  and  herbs  but  they  may 
also  be  found  feeding  upon  the  twigs  and  roots.  They  are  not 
infrequently  attended  by  ants.  The  oviparous  females  are  nearly 
always  apterous,  but  the  males,  on  the  other  hand,  usually  are  winged. 
Apterous  males,  however,  are  common  and  intermediate  forms 
between  alate  males  and  apterous  males  sometimes  occur.  Inter- 


40  BULLETIN   826,   U.    S.   DEPARTMENT   OF   AGRICULTURE. 

mediates  between  apterous  and  alatc  viviparous  forms  are  of  quite 
common  occurrence  and  indicate  possible  origin  of  the  apterous 
forms.  It  is  no  doubt  true,  however,  that  in  members  of  this  tribe 
the  equilibrium  is  disturbed  more  easily  than  in  some  of  the  others, 
and  that  external  influences  have  a  more  sudden  and  noticeable 
effect. 

We  have  divided  the  tribe,  into  four  sub  tribes  which  may  be 
separated  as  follows: 

KEY  TO   THE   SUBTRIBES   OF  THE   APHIDINI. 

1.  Body  covered  with  long  projections CERVAPHIDINA. 

Body  naked  with  the  exception  of  a  few  hairs 2. 

2.  Head  without  prominent  antennal  tubercles APHIDINA. 

Head  with  prominent  antennal  tubercles 3. 

3.  Wings  with  the  radial  sector  normal MACROSIPHINA. 

Wings  with  the  radial  sector  more  or  less  united  with  the  upper  branch  of 

the  media  or  hind  wings  reduced PENTALONINA. 

Subtribe  APHIDINA. 

The  insects  in  the  Aphidina  show  a  great  variation  in  regard  to 
the  cornicles  and  cauda.  Some  have  very  markedly  developed 
cornicles,  others  have,  extremely  small  ones,  while  one  genus  lacks 
them  altogether.  The  cauda  varies  from  very  large  in  genera  like 
Hyalopterus  to  scarcely  any  visible  cauda  in  some  of  the  other 
genera.  Certain  of  the  genera  appear  more  similar  than  others,  for 
example,  Hyalopterus,  Pergandeidia,  and  Brachycolua  all  have  small 
cornicles  and  somewhat  large  caudse.  Certain  other  genera,  while 
appearing  quite  different  in  some  ways,  are  evidently  related. 
Cavariella,  Hyadaphis,  Aspidaphis,  and  Vesiculaphis  all  have  char- 
acters which  are  very  suggestive,  although  there  are  differences 
between  them.  So  also  there  is  a  group  suggesting  Aphis.  The 
various  genera  may  be  separated  as  follows: 

KEY  TO  THE  GENERA  OF  THE  APHIDINA. 

1.  Cornicles  absent Asiphonaphis. 

Cornicles  present 2. 

2.  Cornicles  swollen,  not  sub  cylindrical  or  tapering 3. 

Cornicles  sub  cylindrical  or  tapering  but  sometimes  extremely  short  and 

ringlike 12. 

3.  Fore  wings  with  the  media  once  branched;  apterous  form  with  very  much 

swollen  cornicles Vesiculaphis. 

Fore  wings  with  the  media  twice  branched 4. 

4.  Hind  wings  with  the  cubitus  lacking Carolinaia. 

Hind  wings  with  both  media  and  cubitus  present 5. 

5.  Abdomen  with  a  dorsal  projection  or  tubercle  above  cauda 6. 

Abdomen  without  this  structure , 7. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  41 

6.     Tubercle  very  large,  entirely  covering  posterior  part  of  body;   cornicles 

small,  opening  at  the  side Aspidaphis. 

Tubercle  of  moderate  size,  as  large  as  cauda  in  the  apterous  form ;  cornicles 
normal,  opening  at  the  end Cavariella. 

7.  Cornicles  long,  very  abruptly  and  distinctly  swollen Liosomaphis. 

Cornicles  of  varying  lengths,  gradually  swollen 8. 

8.  Cauda  short  and  abruptly  conical;  cornicles  about  the  same  length 

as  cauda  and  swollen  in  the  middle Brevicoryne. 

Cauda  not  strikingly  short  or  abruptly  conical 9. 

9.  Cornicles  as  short  as  the  width  of  cauda  at  base  or  shorter 10. 

Cornicles  as  long  as  or  longer  than  the  cauda 11. 

10.  Cornicles  minute,  tubercle-like Brachycolus. 

Cornicles  considerably  longer  than  their  diameter;  cauda,  particularly 

in  the  apterous  form,  large  and  long Hyalopterus. 

11.  Cauda  in  the  apterous  form  long  and  broad,  as  long  as  the  cornicles.  .Hyadaphis. 
Cauda    shorter     than     the    cornicles,    not    correspondingly     long    and 

broad Rhopalosiphum. 

12.  Tarsi  atrophied 13. 

Tarsi  normal 14. 

13.  Antennae  of  six  segments;  cornicles  rather  short Atarsos. 

Antennae  of  five  segments;  cornicles  very  long  and  slender Mastopoda. 

14 .  Cauda  apparent!  y  absent,  or  a  mere  rounded  platelike  structure 21. 

Cauda  normal  in  appearance  but  often  very  short. 15. 

15.  Fore  wings  with  media  once  branched 22. 

Fore  wings  with  media  twice  branched 16. 

16.  Antennae  of  five  segments 24. 

Antennae  of  six  segments 17. 

17.  Cornicles  small,  as  short  as  width  of  cauda  at  base,  which  is  without  con- 

striction          18. 

Cornicles  usually  as  long  as  or  longer  than  cauda,  Aphis-like 19. 

18.  Cauda  very  long  and  large  in  the  apterous  form Pergandeidia. 

Cauda  extremely  short  and  subconical Microsiphum. 

19.  Cauda  short  and  abruptly  conical Anuraphis. 

Cauda  elongate  and  constricted  near  base 20. 

20.  Hind  wings  with  media  and  cubitus  present Aphis. 

Hind  winga  with  only  one  oblique  vein Hysteroneura. 

21.  Cornicles  minute,  not  as  long  as  wide Cryptosiphum. 

Cornicles  moderate  in  length Acaudus. 

22.  Antennae  of  six  segments Toxoptera. 

Antenntc  of  five  segments 23. 

23.  Apterous  form  with  a  prominent  median  projection  on  the  vertex  and 

with  four-segmented  antennae;  cornicles  minute Sanbornia. 

Apterous  form  normal Yamataphis. 

24 .  Cornicles  elongate,  Aphis-like Cerosipha. 

Cornicles  extremely  short,  body  of  apterous  form,  much  arched. Siphonatrophia. 

Genus  ACAUDUS  V.  d.  Goot. 

Plate  VI,  A,  B. 

1913.    Acaudus  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  97. 

1917.    Macchiatiella  Del  Guorcio,  Redia,  v.  12,  p.  210. 

1917.    Hannabura  Matsumura,  Journ.  Coll.  Agr.  Tohoku  Univ.,-  v.  7,  pt.  6,  p.  377. 

Van  der  Goot  erected  his  genus  Acaudus  with  lychnidis  L.  as  type. 
This  species  is  quite  similar  to  an  Aphis  without  a  cauda  or  with  a 
very  short,  somewhat  rounded  cauda.  The  cornicles  are  not  long. 


42  BULLETIN   826,   U.   S.   DEPARTMENT  OF   AGRICULTURE. 

Del  Guercio's  gentis  had  his  trifolii  placed  as  type.  The  corni- 
cles of  this  species  appear  to  be  a  little  longer  than  those  of  the  type 
of  Van  der  Goot's  genus,  but  in  other  respects  the  insects  seem  to 
be  quite  similar.  We  feel  that  they  belong  to  the  same  genus.  We 
are  keeping  this  genus  distinct  from  Cryptosiphum,  not  only  because 
of  the  minute  cornicles  in  that  genus  but  also  on  account  of  the 
peculiar  head  structure  which  is  there  seen.  It  is  not  typically  Aphis- 
like. 

CJtaractcrs. — Head  without  prominent  antennal  tubercles.  Antenna?  of  six  seg- 
ments. Fore  wings  with  the  media  twice  branched;  hind  wings  with  both  media 
and  cubitus  present.  Cornicles  cylindrical,  of  moderate  length;  cauda  reduced  to  a 
broad,  short,  rounded  structure. 

Type  (fixed  by  Van  der  Goot,  1913),  Aphis  lychnidis  L. 

Genus  ANURAPHIS  Del  Guerdo. 
Plate  VI,  C-F. 

1907.  Anuraphis  Del  Guercio,  Redia,  v.  4,  p.  190. 

1913.  Brachycaudus  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  97. 

1913.  Dentatiis  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  98. 

1913.  Semiaphis  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  105. 

1917.  Yezabura  Matsumura,  loom.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  392. 

1918.  Sappaphis  Matsumura,  Trans.  Sapporo  Nat.  Hist.  Soc.,  v.  7,  pt.  1,  p.  18. 

In  1907  Del  Guercio  erected  the  genus  Anuraphis  in  which  were 
included  pyri  Koch,  lappae  Koch,  iridis  Del  Guercio,  ranunculi  Kalt., 
myosotidis  Koch,  centaureaeKoch,  prunicola  Kalt.,  tragopogonis  Kalt., 
iani  J?eer,farfarae  Koch,  and  persicae  Boyer. 

These  species,  with  the  exception  of  lappae  and  tragopogonis  which 
seem  to  belong  to  Aphis,  have  a  broadly  and  somewhat  abruptly 
conical  cauda  quite  unlike  that  of  the  genus  Aphis.  In  1913,  Van  der 
Goot  erected  his  genus  Brachycaudus  with  one  of  these  species,  myoso- 
tidis, as  type  without  referring  any  of  the  species  to  Anuraphis  at  all. 
It  is  evident  that  Brachycaudus  is  a  synonym  of  Anuraphis.  At  the 
same  time  he  erected  the  genus  Semiaphis  with  carotae  Koch  as  type. 
A  study  of  this  species,  as  seen  in  Plate  VI,  makes  it  evident  that  this 
is  a  species  of  the  same  general  type.  Semiaphis  thus  becomes  a 
synonym  of  Anuraphis.  Based  on  the  minute  tubercles,  particularly 
on  the  head  and  caudal  portion  of  the  abdomen,  Van  der  Goot  also 
erected  the  genus  Dentatus  with  sorbi  Kalt.  as  type.  Apart  from 
these  tubercles  sorbi  is  in  the  character  of  the  cauda,  etc.,  a  typical 
Anuraphis.  In  our  American  rosy  aphis,  a  species  very  similar  to 
sorbi,  these  tubercles  are  absent  in  many  individuals,  and  in  the  fall 
migrants  the  caudal  ones  are  nearly  always  absent.  If  this  character 
were  retained,  therefore,  the  species  would  belong  in  one  genus  as 
far  as  the  spring  migrant  is  concerned,  and  in  another  genus  when 
the  fall  migrant  is  considered.  Dentatus,  therefore,  becomes  also 
a  synonym  of  Anuraphis. 


GENERIC   CLASSIFICATION   OF  APHIDIDAE.  43 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  six  seg- 
ments and  armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice 
branched,  hind  wings  with  both  media  and  cubitus  present.  Cornicles  cylindrical, 
often  short,  though  sometimes  moderately  long;  catida  short,  broad,  and  abruptly 
conical,  never  elongate  and  constricted  as  in  Aphis.  Males  usually  winged.  Ovi- 
parous forms  apterous. 

Type  (fixed  by  Del  Guercio,  1907),  Aphis  pyri  Koch. 

Genus  APHIS  L. 

Plate  VI,  G-I. 

1758.  Aphis  Linnaeus,  Systems  Naturae,  10th  ed.,  p.  451. 

1817.  Loxerates  Raftnesque,  Am.  Mo.  Mag.  &  Crit.  Review,  v.  1,  p.  361. 

1907.  Uraphis  Del  Guercio,  Redia,  v.  4,  p.  192. 

€  1907.  Microsiphon  Del  Guercio,  Redia,  v.  4,  p.  192. 

1913.  Myzaphis  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  96. 

1913.  Stenaphis  Del  Guercio,  Redia,  v.  9,  p.  185. 

1916.  Longiunguis  Van  der  Goot,  Zur  Kenntniss  der  Blattliiuse  Java's,  p.  112. 

1916.  Melanaphis  Van  der  Goot,  Zur  Kenntniss  der  Blattliiuse  Java's,  p.  61. 

1917.  Afara  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  407. 
1917.  Arimakia  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  405. 

A  number  of  species  were  included  in  the  original  genus  by  Lin- 
naeus. Of  these  Lamarck  set  Aphis  ulmi  L.  as  type  in  1801,  and  in 
1802  Latreille  set  Aphis  sambuci  L.  as  type.  Aphis  ulmi  L. 
is,  according  to  Passerini,  Eriosoma  lanuginosa  Hartig  and  the  spe- 
cies now  placed  in  Tetraneura.  Sambuci  is  retained  here  as  type 
and  a  request  will  be  submitted  that  this  species  be  fixed  definitely 
by  the  International  Commission.  It  would  greatly  disarrange  the 
economic  literature  to  change  the  meaning  of  this  common  name. 

In  1907  Del  Guercio  erected  two  genera,  Uraphis  and  Microsiphon 
based  on  th^p  relative  length  of  the  cornicles  and  cauda.  When  con- 
sidering certain  individual  species  this  would  appear  as  a  very  fair 
character  for  use.  But  when  large  series  of  species  are  studied  it  will 
be  found  that  in  the  species  having  the  cauda  of  the  typical  Aphis 
shape  there  are  all  gradations  of  cornicles  from  the  very  short  to 
the  very  long.  This  will  be  seen  also  in  the  forms  having  the  abruptly 
conical  cauda.  Some  species  have  very  short  cornicles  and  some 
quite  long  ones.  Moreover,  in  the  same  species  the  cornicles  in  the 
different  forms  will  bear  a  different  relation  to  the  length  of  the 
cauda.  Species,  therefore,  having  cornicles  and  cauda  of  essentially 
the  same  character  should  not  be  used  as  types  of  different  genera 
depending  on  the  length  of  the  cornicles.  If  this  were  the  case, 
certain  species  which  are  close  to  the  border  line  of  separation  would 
on  some  individuals  fall  in  one  genus  and  on  other  individuals  fall  in  a 
different  genus.  Under  this  rule,  saccliari  Zehnt.  would,  it  is  believed, 
belong  to  the  genus  Aphis,  and  Longiunguis,  therefore,  be  a  synonym 
of  Aphis.  Likewise  the  genus  Melanaphis  would  be  a  synonym  of 
Aphis.  This  genus  was  erected  with  bambusae  Kirk,  as  type.  The 
cornicles  are  short  but  the  cauda  is  aphis-like.  Where  the  generic 
characters  are  considered,  the  genus  appears  a  synonym. 


44  BULLETIN   826,   TJ.   S.   DEPARTMENT   OF   AGRICULTURE. 

The  genus  Myzoaphis  was  erected  by  Van  der  Goot  with  Aphis 
rosarum  Kalt.  and  Aphis  abietina  Walker.  These  two  species  show 
practically  no  antennal  tubercles  and  are  very  little  different  from 
a  typical  Aphis,  excepting  in  the  clothing  of  the  antennae  and  body. 
The  cauda  as  figured  by  Van  der  Goot  is  quite  conical,  but  the 
writer's  specimens  are  somewhat  different  from  this  and  specimens 
of  abietinus  do  not  show  a  cauda  exactly  like  his  figure.  In  fact, 
they  appear  more  like  an  Aphis.  This  genus,  therefore,  should  not 
be  separated  or  it  will  necessitate  the  separation  of  very  many 
other  forms  under  new  names. 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  six  seg- 
ments and  armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice  branched ; 
hind  wings  with  both  media  and  cubitus  present.  Cornicles  cylindrical  or  slightly 
tapering.  Cauda  usually  not  as  long  as  the  cornicles,  subconical,  rather  elongate, 
constricted  about  the  middle.  Anal  plate  rounded.  Males  usually  winged,  oviparous 
females  apterous. 

Type  (by  suspension  of  rules),  Aphis  sambuci  L. 

Genus  ASPIDAPHIS  Gillette. 

Plate  VI,  L-O. 
1917.    Aspidaphis  Gillette,  Can.  Ent.,  v.  49,  p.  196. 

The  genus  Aspidaphis  appears  to  be  related  both  to  Cavariella 
and  to  Vesiculaphis.  It  has  the  short,  blocky  form  of  Vesiculaphis 
and  also  the  peculiar  integument.  On  the  other  hand,  certain  species 
with  a  similar  integument  are  met  with  in  Cavariella.  The  develop- 
ment of  the  dorsal  abdominal  tubercle  is  here  very  pronounced  and 
the  cornicles  have  taken  on  a  peculiar  shape.  „ 

Characters. — Head  without  prominent  antennal  tubercles;  antennae  short,  of  five 
segments,  armed  with  subcircular  sensoria.  Wing  venation  normal.  Cornicles  very 
small,  somewhat  swollen  near  the  distal  extremity  and  with  the  opening  in  the  side  of 
the  cornicle,  not  at  the  tip.  Abdomen  with  a  dorsal  caudal  tubercle  developed  into 
a  large  conical  process  extending  beyond  and  fully  covering  the  cauda  in  the  apterous 
form.  Body  elongate. 

Type  (monotypical),  Aspidaphis  polygonii  Gill. 

Genus  ASIPHONAPHIS  Wilson  &  Davis. 
1919.  AsiphonapJiis  Wilson  &  Davis,  Ent.  News,  v.  30,  p.  39. 

Characters.— Head  without  prominent  antennal  tubercles.  Antennae  of  six  segments 
armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice  branched;  hind 
wings  with  both  media  and  cubitus  present.  Cornicles  absent  entirely.  Abdomen 
with  large  lateral  tubercles.  Cauda  somewhat  conical  or  Aphis^like.  Anal  plate 

rounded. 

Type  (monotypical),  Asiphonaphis  pruni  Wilson  &  Davis. 

Genus  ATARSOS  Gillette. 

Plate  VI,  P-S. 

1911.    Atarsos  Gillette,  Ent.  News,  v.  22,  p.  440. 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  six  segments 
armed  with  subcircular,  somewhat  tuberculate  sensoria;  fore  wings  with  the  media 
twice  branched,  hind  wings  with  both  media  and  cubitus  present.  Cornicles  rather 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  45 

short,  subcylindrical ;  cauda  somewhat  conical,  not  quite  as  long  as  the  cornicles;  anal 
plate  rounded.     Tarsi  atrophied  in  all  the  forms. 

Type  (monotypical),  Atarsos  grindeliae  Gill. 

Genus  BRACHYCOLUS  Buckton. 
Plate  VI,  T,  U. 

1879.    Brachycolus  Buckton,  Mon.  British  Aphides,  v.  2,  p.  146. 
1913.    Brachysiphum  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  105. 

The  genus  Brachycolus  was  erected  with  stellariae  Hardy  as  type 
and  is  easily  recognized  from  the  structure  of  the  cornicles.  Van 
der  Goot's  genus  is  in  all  essential  respects  the  same,  the  type  of  that 
genus  being  fhalictri  Koch. 

Cftaracters. — Head  without  prominent  antennal  tubercles.  Antennee  of  six  segments 
and  armed  with  subcircular  sensoria.  Wing  venation  normal.  Cornicles  very  small, 
especially  in  the  apterous  form.  Cauda  medium  in  size  and  conical.  Body  elongate; 
legs  and  antennae  usually  short. 

Type  (monotypical),  Aphis  stellariae  Hardy. 

Genus  BREVICORYNE  V.  d.  Goot. 
Plate  VI,  J,  K. 

1915.  Brevicoryne  Van  der  Goot,  Beitrage  z.  Kennt.  d.  Holl.  Blattliiuse,  p.  245. 

1916.  Oedisiphum  Van  der  Goot,  Zur  Kenntniss  der  Blattliiuse  Java's,  p.  122. 
1918.    Brevicoryne  Das,  Mem.  Ind.  Mus.,  v.  6,  p.  179. 

The  genus  Brevicoryne  was  erected  with  Aphis  Irassicae  L.  as  type, 
a  species  in  which  the  cornicles  are  very  short  and  somewhat  swollen 
in  the  middle,  and  the  cauda  conical.  In  the  species  Oedisiphum 
cojnpositarum  V.  d.  Goot  quite  similar  characters  are  found.  The 
cornicles  appear  to  be  somewhat  more  slender  but  in  the  main  the 
species  appear  alike.  Oedisiphum,  therefore,  becomes  a  synonym. 

Characters. — Head  without  prominent  antennal  tubercles;  antennae  of  six  segments 
and  armed  with  subcircular  sensoria.  Wing  venation  normal.  Cornicles  short,  not 
much  longer  than  the  cauda  and  swollen  in  the  middle.  Cauda  short  and  broadly 
conical. 

Forms  not  especially  elongate. 

Type  (monotypical),  Aphis  brassicae  L. 

Genus  CAROLINAIA  Wilson. 

Plate  VI,  V,  \V. 
1911.     Carolinaia  Wilson,  Can.  Ent.,  v.  43,  p.  61. 

The  genus  Carolinaia  Wilson  is  related  very  closely  to  Rhopalosi- 
phum.  In  fact  some  of  the  species  placed  here  are  distinguished 
from  members  of  that  genus  only  by  the  fact  that  the  cubitus  is 
lacking  in  the  hind  wing.  It  is  true  that  the  type  species  was 
described  as  having  five-segmented  antennae  in  the  apterous  form. 
But  other  species,  like  this  in  all  other  respects,  have  normal  six- 
segmented  antennas.  Certain  species  of  Rhopalosiphum  will  show 
strains  in  which  nearly  all  of  the  individuals  will  have  five-segmented 
antennae  and  yet  the  normal"  antenna  for  such  species  is  a  six- 
segmented  one.  In  this  closely  related  genus,  therefore,  it  is  not 
surprising  if  a  similar  condition  is  met  with. 


46  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  five  or  six 
segments  armed  with  subcircular  sensoria.  Venation  of  the  fore  wings  normal;  hind 
wings  with  the  cubitus  absent;  cornicles  elongate,  slightly  swollen  near  the  distal 
extremity,  but  without  a  prominent  neck  near  the  proximal  end.  Cauda  rather 
broadly  conical. 

Type  (monotypical),  Carolinaia  caricis  Wlsn. 

Genus  CAVARIELLA  Del  Guercio. 
Plate  VI,  X-Z. 

1911.     Cavariella  Del  Guercio,  Redia,  v.  7,  p.  323. 

1914.     Corynaziphon  Mordwilko,  Faune  de  la  Russie  Insecta,  Aphidodea,  p.  73. 

1917.  Nipposiphum  Matsumura,  Journ.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  410. 

Two  new  genera  were  erected  in  recent  years:  One  by  Del 
Guercio,  Cavariella,  with  pastinacae  L.  as  type;  the  other  by 
Mordwilko,  called  Corynosiphon,  but  with  no  species  definitely 
given.  In  a  footnote,  however,  Mordwilko  refers  capre&e  Fab.  to 
his  Corynosiphon.  Both  of  these  genera,  therefore,  were  used  for 
insects  of  the  same  general  type.  This  may  be  said  too  of 
Nipposiphum. 

Characters. — Head  without  distinct  antennal  tubercles;  antennae  of  six  segments, 
armed  with  prominent  sensoria.  Wing  venation  normal.  Cornicles  somewhat  swollen 
near  the  distal  end;  cauda  rather  elongate,  somewhat  conical;  abdomen  with  a  tubercle 
or  horn  above  the  cauda,  this  tubercle  or  horn  most  prominent  in  the  apterous  form. 
Males  winged;  oviparous  females  usually  apterous. 

Type,  Aphis  pastinacae  L.  1 

Genus  CEROSIPHA  De^Guercio. 

1900.     Cerosipha  Del  Guercio,  Nuove  Rel.  Staz.  Firenze,  ser.  1,  no.  2,  p.  116. 

1918.  Metaphis  Matsumura,  Trans.  Sapporo  Nat.  Hist.  Soc.,  v.  7,  pt.  1,  p.  1. 

The  genus  Cerosipha  Del  Guercio  was  erected  for  a  species,  pas- 
seriniana  Del  Guercio,  somewhat  similar  to  Aphis  but  with  five  seg- 
ments to  the  antennae.  The  writer  has  been  unable  to  study  the  type 
species  and  has  based  his  remarks  on  rubifolii  Thomas,  an  American 
species. 

Characters. — Head  without  prominent  antennal  tubercles;  antennae  of  five  segments, 
Aphis-like,  armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice 
branched,  hind  wings  with  both  media  and  cubitus  present.  Cornicles  cylindrical  or 
somewhat  tapering,  cauda  Aphis-like,  somewhat  tapering. 

Type  (monotypical),  Cerosipha  passeriniana  Del  G. 

Genus  CRYPTOSIPHUM  Buckton. 

Plate  VI,  PP,  QQ. 
1879.     Cryptosiphum  Buckton,  Hon.  Br.  Aphides,  v.  2,  p.  144. 

The  genus  Cryptosiphum  Buckton  can  be  distinguished  from  others 
having  very  short  cornicles  by  the  cauda  which  is  here  short  and 
rounded,  while  in  most  other  cases  it  is  very  long. 

Characters. — Head  without  distinct  antennal  tubercles.  Antennae  of  six  segments, 
rather  short,  and  with  subcircular  sensoria.  Fore  wings  with  the  media  either  once  or 
twice  branched.  Hind  wings  with  both  media. and  cubitus  present.  Cornicles  sub-. 

1  The  writer  has  a  record  to  the  eSect  that  in  a  published  paper  Del  Guercio  used  pastinacae  as  type 
of  Cavariella,  but  he  has  been  unable  to  locate  the  publication  and  can  not  reach  Doctor  Del  Guercio 
through  the  mails. 


GENEKIC   CLASSIFICATION   OF  APHIDIDAE.  47 

Cylindrical  but  extremely  short,  not  as  long  as  wide.     Cauda  very-  short  and  rounded, 
not  Aphis-like;  anal  plate  rounded. 

Type  (monotypical),  Cryptosiphum  artemisiae  Buckt. 

Genus  HYSTERONEURA  Davis. 

1919.     Heteroneura  Davis,  Can.  Ent.,  v.  51,  p.  228. 
1919.    Hysteroneura  Davis,  Can.  Ent.,  v.  51,  p.  263. 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  six  seg- 
ments armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice  branched. 
Hind  wings  with  the  cubitus  absent.  Cornicles  somewhat  tapering  or  eubcylindric, 
cauda  Aphis-like;  anal  plate  rounded. 

Type  (monotypical),  Aphis  setariae  Thos. 
Genus  HYADAPHIS  Kirk. 
Plate  VI,  AA,  BB. 

1904.     Hyadaphis  Kirkaldy,  The  Entomologist,  v.  37,  p.  279. 

1863.    Siphocoryne  Passerini,  Aphididse  Italicse,  p.  8  (not  Siphocoryne  ,  1860). 

As  indicated  under  the  discussion  of  Siphocoryne,  Passerini  set 
xylostei  as  type  of  his  genus  in  1863.  Nymphaeae  had,  however, 
been  set  in  1860,  so  Kirkaldy  gave  Hyadaphis  to  Passerini 's  1863 
conception,  of  which  genus  xylostei  becomes  the  type.  Xylostei  has  no 
caudal  horn  and  is  quite  similar  in  general  appearance  to  a  Rhopalo- 
siphum.  We  may  separate  the  two  genera,  however,  on  the  cauda, 
which  in  xylostei,  particularly  in  the  apterous  form,  is  very  large,  fully 
as  long  as  the  cornicles,  and  broad,  quite  unlike  that  of  nymphaeae- 
Several  other  species  which  have  generally  been  considered  in  the 
same  genus  with  xylostei  possess  a  distinct  caudal  projection  on  the 
abdomen.  One  of  these  has  been  made  the  type  of  Cavariella  so  that 
such  species  will  be  removed  from  our  conception  of  Hyadaphis. 

Characters. — Head  without  prominent  antennal  tubercles;  antennae  of  six  segments 
which  are  rather  abundantly  armed  with  tuberoulate  sensoria.  Wing  venation  normal. 
Cornicles  somewhat  swollen  but  not  prominently  so.  Cauda,  particularly  in  the 
apterous  form,  large,  as  long  as  the  cornicles,  and  broad.  Males  usually  winged; 
oviparous  females  apterous;  summer  forms  usually  feeding  on  the  TJmbelliferae. 

Type  (fixed  by  Kirkaldy,  1904),  Aphis  xylostei  Schrk. 

Genus  HYALOPTERUS  Koch. 
Plate  VI,  RR-WW. 

1854.     Hyalopterus  Koch,  Die  Fflanz.  Aphiden,  p.  10. 
1917.     Hayhurstia  Del  Ouercio,  Rcdla,  v.  12,  p.  208. 

Aphis  pruni  Fab.  was  set  as  the  type  of  this  gonus  in  1860  by 
Passerini.  In  1917  Del  Guercio  erected  the  genus  Haylmrstia. 
There  is  little  difference  between  the  two  as  will  be  seen  by  examining 
the  drawings  given  herewith.  The  cauda  in  Hayhurstia  is  slightly 
narrower  than  that  in  Hyalopterus.  It  is  the  writer's  opinion  that 
the  two  represent  only  one  genus. 

Two  other  genera  may  be  mentioned  here.  They  are  Brachycolus 
Buckt.  and  Pergaiideidia  Schout.  Specimens  of  the  type  species  of 
Brachycolus  show  that  this  genus  is  quite  similar  to  Hyalopterus, 
but  it  differs  in  the  cornicles.  In  Brachycolus  the  cornicles  are  very 
small,  almost  mere  rounded  swellings  in  the  apterous  form,  whereas 


48  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

in  Hyalopterus  the  cornicles  are  in  the  apterous  form  of  fair  size  and 
not  distinctly  swollen. 

Specimens  of  the  type  of  Pergandeidia  received  from  Schouteden 
show  that  this  genus  is  very  close  to  Hyalopterus  and  is  probably 
almost  too  close  for  a  very  distinct  genus.  The  diagnosis  given  by 
Wilson  (1910)  for  this  genus  does  not  agree  with  the  type  species  as 
determined  by  the  author  of  the  genus. 

In  the  specimens  examined  by  the  writer  the  cauda,  as  will  be  seen 
in  the  drawings,  is  very  much  longer  than  the  cornicles,  bearing  about 
the  same  ratio  as  seen  in  Hyalopterus. 

Characters. — Head  without  distinct  antennal  tubercles,  antennae  of  six  segments 
armed  with  subcircular  sensoria.  Wing  venation  normal.  Cornicles  very  short,  not 
much  longer  than  the  cauda  is  wide  at  its  base,  swollen  beyond  the  middle,  particularly 
in  the  alate  form.  Cauda  long  and  broad,  considerably  longer  than  the  cornicles. 
Form  of  the  insects  elongate,  often  more  or  less  flat. 

Type  (fixed  by  Passerini,  1860),  Aphis  pruni  Fab.  (=A.  arundinis  Fab.). 

Genus  LIOSOMAPHIS  Walker. 

Plate  VI,  NN,  OO. 
1868.    Liosomaphis  Walker,  The  Zoologist,  p.  1119. 

The  genus  Liosomaphis  Walker  is  related  somewhat  closely  to 
Khopalosiphum.  The  two  genera,  however,  can  be  separated  on  the 
structure  of  the  cornicles. 

In  Liosomaphis  the  cornicles  have  a  very  distinct  neck  near  the 
proximal  extremity,  due  to  a  constriction  behind  the  prominent 
swelling.  This  is  strikingly  evident  in  the  apterous  form,  as  well  as 
in  the  alate  one.  In  Rhopalosiphum,  on  the  other  hand,  there  is  no 
ab-iupt  swelling,  but  only  a  gradual  one  which  is  not  at  all  prominent, 
as  in  Liosomaphis. 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  six  segments 
armed  with  subcircular  sensoria.  Wing  venation  normal.  Cornicles  elongate, 
distinctly  swollen  in  the  middle,  and  with  a  constricted  neck  near  the  base. 
Structure  in  both  the  apterous  and  alate  forms  similar.  Cauda  not  as  long  as  the 
cornicles,  somewhat  narrowly  conical.  Males  usually  winged;  oviparous  females 
usually  apterous. 

Type  (monotypical),  Aphis  berberidis  Kalt. 

Genus  MASTOPODA  Oestlund. 
1886.     J/astop<x2a.Oestlund,  Minn.  Geol.  Surv.  Kept.  14,  p.  52. 

The  genus  Mastopoda  Oestlund,  like  Gillette's  Atarsos,  is  peculiar 
in  that  the  tarsi  are  atrophied. 

Characters. — Head  without  distinct  autennal  tubercles.  Antennae  of  five  segments. 
Fore  wings  with  the  media  twice  branched,  hind  wings  with  both  media  and  cubitus 
present.  Cornicles  somewhat  long  and  cylindrical.  Cauda  short,  conical,  Aphis- 
like.  Legs  with  the  tarsi  absent  and  provided  instead  with  a  membranous  disk 
which  enables  the  insect  to  walk  inverted  on  smooth  surfaces. 

Type  (monotypical),  Mastopoda  pteridis  Oestlund. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  49 

Genus  MICROSIPHUM  Chotodkovsky. 
1908.     Microsiphum  Cholodkovsky,  Zool.  Anz.,  v.  32,  p.  687. 

Characters. — Head  without  prominent  antenna!  tubercles,  although  with  apparent 
ones.  Antennae  of  six  segments,  rather  long  and  slender.  Fore  wings  with  the  media 
twice  branched;  hind  wings  with  both  media  and  cubitus  present.  Cornicles  very 
short,  not  much  longer  than  wide  and  often  scarcely  visible  in  the  alate  form.  Cauda 
extremely  short  and  subconical  or  slightly  rounded;  anal  plate  rounded. 

Type  (monotypical),  Microsiphum  ptannicae  Choi. 

Genus  PERGANDEIDIA  Schouteden. 
Plate  VI,  KK-MM. 

1903.  Pergandcidia  Schouteden,  Zool.  Anz.,  v.  2fi,  p.  CSS. 

1913.  Longicaudus  Van  der  Goot,  Tijd.  voor  Ent.,  v.  5G,  p.  140. 

1915.  Rhizobcrlesia  Del  Guercio,  Redia,  v.  10,  p.  240. 

1918.  Yezosiphum  Matsumura,  Trans.  Sapporo  Nat.  Hist.  Soc.,  v.  7,  pt.  1,  p.  7. 

1918.  Brachyunguis  Das,  Mem.  Ind.  Mus.,  v.  6,  p.  227. 

The  genus  Pergandeidia  was  erected  by  Schouteden  with  his 
ononidis  as  type.  The  characterization  of  the  genus  in  this  paper  is 
based  on  specimens  of  the  species  received  from  Schouteden,  and 
though  it  does  not  agree  with  the  characterization  of  the  genus  as 
sometimes  given,  it  seems  necessary  to  follow  the  type  species  in 
forming  a  conception  of  the  genus.  In  1913  Van  der  Goot  erected 
the  genus  Longicaudus  and  placed  trirhodus  Walker  as  type.  In 
studying  this  species  differences  sufficient  for  good  generic  distinc- 
tion have  not  been  found,  and,  therefore,  the  conclusion  may  be 
drawn  that  Longicaudus  is  a  synonym  of  Pergandeidia.  Del  Guercio 's 
genus  is  placed  here  also,  although  a  study  of  the  type  species,  which 
is  not  available,  possibly  may  show  the  cauda  to  be  somewhat  different. 

Characters. — Head  with  no  prominent  antennal  tubercles;  antennae  of  six  segments. 
Fore  winga  with  media  twice  branched,  hind  wings  with  both  media  and  cubitus 
present.  Cornicles  very  short,  almost  as  wide  as  long.  Cauda  very  long  and  broad. 

Type  (monotypical),  Pergandeidia  ononidis  Schout. 

Genus  RHOPALOSIPHUM  Koch. 

Plate  VI,  FF-JJ. 

1-854.  Rhopalosiphum  Koch,  Die  Tflanz.  Aphidcn,  p.  23. 

1860.  Siphocoryne  Passerini,  Gli  Afuli,  p.  28. 

1882.  Rhopalosiphon  Scudder,  Nomenclator  Zoologicus. 

1910.  Coloradoa  Wilson,  Ann.  Ent.  Soc.  Am.,  v.  3,  p.  323. 

1915.  Siphonaphis  Van  der  Goot,  Beitriige  zur  Kennt.  d.  Holl.  Blattlause,  p.  238. 

1918.  Stcphcmonia  Das,  Mem.  Ind.  MILS.,  v.  e,  p.  175. 

1856.  (Type  fixation)  Gerstaeckcr,  Bericht  for  1854,  p.  162. 

There  has  been  much  confusion  in  regard  to  this  genus.  Koch 
included  a  number  of  diverse  species.  In  1856  Gerstaecker  defi- 
nitely set  Aphis  nymplieae  L.  as  type.  Overlooking  this,  Passerini 
in  1860  set  persicae  Sulz.  as  type.  In  1863,  however,  he  changed  this 
name  to  diantlii  Schrank.  In  1860  Passerini  erected  the  genus  Sipho- 
coryne with  A.  nymplieae  L.  as  type.  Having  the  same  type  this 
must,  therefore,  become  a  synonym  of  Rhopalosiphum.  In  1863 
Passerini  placed  nymplieae  in  Rhopalosiphum,  although  he  set  diantlii 
141613°—  20— Bull.  826 4 


50  BULLETIN    826,    U.   S.    DEPARTMENT   OF   AGRICULTURE. 

as  type,  and  he  used  Siphocoryne  in  a  different  sense  with  xylostei 
Schrank  as  type.  Such  a  procedure  is  inadmissible,  and  Kirkaldy 
therefore  renamed  this  genus  Hyad  aphis. 

The  genus  Coloradoa  Wilson  was  erected  with  rufornaculata  Wilson 
as  type.  Although  much  smaller,  this  species  is  essentially  like 
nympTieae  in  structure,  and  we  therefore  consider  the  genus  a  syno- 
nym of  Rhopalosiphum. 

The  genus  Siphonaphis  Van  der  Goot  was  erected  with  nympJieae 
L.  as  type.  Having  the  same  type,  therefore,  it  must  become  a 
synonym  of  Rhopalosiphum.  Lahorensis  Das  is  quite  similar. 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  six  seg- 
ments with  the  usual  subcircular  sensoria  present.  Wing  venation  normal.  Cor- 
nicles moderately  long  and  slender,  slightly  swollen  near  their  distal  extremities. 
Cauda  rather  elongate,  not  as  long  as  the  cornicles,  and  not  broad.  Abdomen  without 
a  caudal  horn  or  projection  above  the  cauda.  Males  usually  winged;  oviparous 
females  usually  apterous. 

Type  (fixed  by  Gerstaecker,  1856),  Aphis  nympheae  L. 
Genus  SANBORNIA,  n.  gen. 
Plate  VII,  F-L. 

The  genus  Sanbornia  is  erected  for  a  peculiar  form  living  on 
juniper  at  College  Station,  Tex.,  and  forwarded  to  the  Bureau  by 
Charles  Sanborn.  This  species  was  determined  as  undescribed  by  Mr. 
Pergande  who  had  planned  to  publish  on  it.  He  had  given  it  the 
name  juniperi.  The  type  is  in  the  U.  S.  National  Museum  (Cat. 
No.  23064). 

Characters. — Head  without  prominent  antennal  tubercles.  Antennae  of  five  seg- 
ments, armed  with  circular  sensoria.  Fore  wings  with  the  media  once  branched, 
hind  wings  with  only  the  media  present,  cornicles  minute;  cauda  elongate.  Apterous 
form  with  four-segmented  antennae  and  with  a  prominent  mushroom-like  projection 
on  the  vertex. 

Type,  Sanbornia  juniperi  Perg. 

Sanbornia  juniperi  Pergande,  n.  sp. 
(Description  by  Mr.  T.  Pergande.) 

Apterous  form. — The  head  is  most  remarkable  in  front,  having  a  large,  squarish, 
bilobed  projection  about  the  middle  and  each  side  of  it;  close  to  the  insertion  of  the 
antennse  is  a  prominent,  short,  and  conical  protuberance.  There  is  also  at,  the  inner 
side  of  the  first  antennal  joint  a  long  and  slightly  conical  protuberance.  The  antennae 
are  but  four-jointed,  the  spur  shorter  than  the  basal  section  of  the  joint.  Nectaries 
are  not  visible  (?);  the  tail  is  rather  long  and  uniformly  elongate  conical;  the  tarsi 
are  very  short,  the  first  joint  appears  to  be  minute,  and  in  alcoholic  specimens  seems 
to  be  withdrawn  into  the  tibiae;  the  last  abdominal  segment  is  semicircular. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  51 

Genus  SIPHONATROPHIA  Swain. 
Plate  VII,  A-E. 

1918.    SiphonatropJiia  Swain,  Ent.  News,  v.  29,  p.  363. 

Characters. — Head  without  prominent  antenna!  tubercles.  Antennae  of  five  seg- 
ments and  armed  with  circular  sensoria.  Fore  wings  with  the  media  twice  branched; 
hind  wings  with  both  media  and  ctfbitus  present.  Cauda  elongate  and  conical.  Cor- 
nicles extremely  short.  Body  of  apterous  form  considerably  arched. 

Type  (monotypical),  .Cerosipha  cupressi  Swain. 

Genus  YAMATAPHIS  Matsumura. 

1917.     Yamataphis  Matsumura,  Jour.  Coll.  Agr.  Tohoku  Univ.,  v.  7,  pt.  6,  p.  412. 
Characters. — Head  without  prominent  antennal  tubercles.     Antennae  of  five  seg- 
ments armed  with  small  circular  sensoria.    Fore  wings  with  the  media  once  branched; 
hind  wings  with  both  media  and  cubitus  present.     Cornicles  subcylindrical.     Cauda 
Somewhat  conical. 

Type  (fixed  by  Matsumura,  1917),   Yamataphis   oryzae  Mats. 

Genus  TOXOPTERA  Koch. 
Plate  VI,  DD,  EE. 

1857.     Toxoptcra  Koch,  Die  Pflanz.  Aphidcn,  p.  253. 
1891.     Ceylonia  Buckton,  Ind.  Mus.  Notes,  v.  2,  p.  35. 

Koch's  genus  was  erected  for  his  aurantiae,  a  well-known  species  on 
citrus,  etc.  Buckton  erected  his  genus  for  a  species  he  described  as 
tlieaecola.  Specimens  of  aurantiae  from  various  regions  and  specimens 
of  tlieaecola  from  Zehntner  substantiate  the  placing  of  theaecola  as  a 
synonym  of  aurantiae.  Ceylonia  will  then  become  a  synonym  of 
Toxoptera.  Even  if  the  two  species  were  held  to  be  distinct,  this 
would  necessarily  be  the  case. 

diameters. — Head  without  prominent  antennal   tubercles.     Antennae  of   six  seg- 
ments, armed  \vith  subcircular  sensoria.     Fore  wings  with  the  media  once  branched, 
hind  wings  with  both  media  and  cubitus  present.     Cornicles  moderate  in  length,  sub- 
cylindric,  tapering.     Cauda  of  moderate  length,  somewhat  constricted  near  the  base. 
Type  (monotypical),  Toxoptera  aurantiae  Koch  (aurantiae  Boyer). 

Genus  VESICULAPHIS  Del  Guercio. 

Plate  VII,  M-Q. 

1911.     Vesiculaphis  Del  Guercio,  Redia,  v.  7,  p.  4(54. 

Characters. — Apterous  form  elongate ;  antennae  short,  composed  of  five  segments  and 
situated  on  the  under  side  of  the  head.  Top  of  head  forming  a  ledge  which  extends 
out  over  the  antennae  forming  an  angle  in  front  of  the  eye;  eyes  protruding.  Cornicles 
large,  very  much  swollen  and  curved,  opening  minute  and  flanged;  cauda  somewhat 
conical,  rounded  at  the  tip.  Anal  plate  rounded;  posterior  part  of  abdomen  extending 
out  over  the  anal  plate  and  somewhat  over  the  cauda.  Alate  form  with  six-segmented 
antenme  armed  with  subcircular  tuberculate  sensoria.  Fore  wings  with  the  media 
once  branched,  hind  wings  with  both  media  and  cubitus  present.  Cornicles  somewhat 
slender,  swollen,  and  slightly  constricted  at  the  tip.  Cauda  conical,  not  as  long  as 
the  cornicles,  anal  plate  rounded. 

Type  (monotypical),  Toxoptera  carlcis  Fullaway. 

This  genus  is  one  of  those  peculiar  genera,  like  Aspidaphis  Gillette, 
which  seem  to  be  related,  to  Rhopalosiphum. 


52  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Subtribe  CERVAPHIDINA. 

The  subtribe  Cervaphidina  is  a  most  interesting  and  peculiar  one. 
The  specialization  is  remarkable  in  that  long  processes  are  developed 
on  the  body  and  considerable  reduction  has  taken  place  in  the  wings 
and  antennae  while  no  Aphis-like  cauda  is  found.  Only  two  genera 
are  known,  which  may  be  separated  as  follows: 

KEY  TO  THE  GENERA  OF  THE  CERVAPHIDINA. 

Cornicles  swollen;  body  spine-like  projections  not  armed  with  teeth Anomalaphis. 

Cornicles  not  swollen;  body  spine-like  projections  armed  with  teeth Cervaphis. 

Genus  ANOMALAPHIS,  n.  gen. 
Plato  VIII,  D-F. 

Characters. — Body  armed  with  elongate  tubercle-like  projections,  particularly  on 
the  caudal  portion;  antennae  five-segmented  in  both  apterous  and  alate  forms, 
armed  with  subcircular  sensoria.  Fore  wings  with  the  media  once  branched; 
hind  wings  considerably  reduced,  with  the  cubitus  absent.  Cornicles  distinctly 
swollen;  cauda  and  anal  plate  reduced. 

Type  Anomalaphis  comperei  Pergande. 

Anomalaphis  comperei  Pergande,  n.  sp. 

Among  the  many  descriptive  notes  left  by  Mr.  Theo.  Pergande  are 
some  recording  a  peculiar  species  from  Australia.  This  proves  to 
represent  an  undescribed  genus  in  the  Cervaphidina.  Pergande  rec- 
ognized the  species  as  typical  of  a  new  genus  to  which  he  gave  the 
manuscript  name  here  used.  He  left  no  description  of  the  genus 
and  his  notes  on  the  species  are  given  here  exactly  as  he  left  them. 
The  type  is  in  the  United  States  National  Museum  collection  of 
Aphididae  (Cat,  No.  23065). 

Feb.  18,  1907,  Rec.  from  Compere,  a  lot  of  Aphides,  found  in  1901,  on  Acacia  and 
Eucalyptus,  along  the  beach  at  Albany,  West  Australia,  which  represents  a  new  genus 
among  Rhopalosiphins,  and  is  a  most  remarkable  Aphid  in  various  respects.  The 
antenna,  in  the  apterous  and  migratory  female,  are  but  5-jted.  while  the  spur  is  rather 
short  and  resembles  that' of  Chaitophorus.  The  front  wings  are  ample  and  reach  con- 
siderably beyond  the  end  of  the  body,  with  the  third  discoidal  having  but  one  fork, 
as  in  Schizoneura.  The  hind  wings  are  very  short  and  narrow  and  reach  out  to  the 
apex  of  the  1st  vein  of  the  anterior  wings,  there  is  also  but  1  discoidal,  straight,  and 
near  the  apex  of  the  wing.  The  nectaries  are  short,  clavate,  and  similar  to  those  of 
Siphocoryne,  the  tail  appears  to  be  wanting.  The  abdomen  of  the  migrant  appears 
to  have  been  of  a  dusky  yellowish  green,  with  transverse  rows  of  small,  black  spots  or 
tubercles,  and  blackish  sutures  between  the  segments.  The  eyes  are  brown;  antennae 
black,  rather  short,  reaching  barely  to  the  abdomen  and  but  5-jointed;  the  two  basal 
joints  as  usually;  the  3rd  joint  is  longest,  about  as  long  as  the  remaining  joints  together, 
including  the  spur,  with  some  projecting  sensoria  and  a  few  short  hairs;  joints  4  and  5 
are  subequal  in  length,  exclusive  of  the  spur,  and  clavate,  the  spur  is  about  $  the 
length  of  the  basal  section  of  the  joint,  rather  stout  and  blunt;  the  front  of  the 
head  resembles  that  of  ApMs.  The  sides  of  the  abdominal  segments  are  somewhat 
angulated,  each  angle  provided  with  a  very  short,  capitate,  stout  bristle,  while  at  the 
posterior  edge  of  the  two  segments,  following  the  nectaries,  there  is  a  pair  of  long, 


GENERIC    CLASSIFICATION    OF   APHIDIDAE.  53 

diverging,  fleshy  spines,  with  a  sharp,  quite  long  and  slender  spine  at  the  tip;  the 
posterior  pair  longest;  all  of  them  black;  the  end  of  the  body  is  fringed  with  fine  and 
quite  long  hairs.  Legs  as  in  other  Aphides. 

The  apterous  females  are  dark  brownish  or  grayish  green  above,  with  a  somewhat  fusi- 
form median,  yellowish  strip,  broadest  near  the  head,  tapering  posteriorly  to  a  point 
and  terminating  in  front  of  nectaries;  the  sutures  of  the  segments,  the  sides  and  under 
side  of  the  body  are  also  of  a  yellowish  color,  on  account  of  which,  there  is  each  side  a 
subdorsal  row  of  transverse,  dark  spots.  The  head  and  about  basal  half  of  the  antennae, 
dark,  dirty  yellowish,  the  eyea  dark  brown.  There  are  about  4  short  and  curved 
capitate  hairs  on  the  front  of  the  head  and  prominent  fleshy  tubercles  each  side  of  the 
body,  each  bearing  at  its  apex  a  short,  capitate  spine  or  hair,  all  of  them  growing  longer 
toward  the  nectaries,  while  beyond  the  nectaries  there  are  two  pairs  of  long  and  slender 
fleshy  tubercles,  tipped  with  a  spine,  ?s  in  the  migrant.  A  tail  could  not  be  seen.  In 
the  younger  forms  and  pupae,  the  tubercles  are  as  in  the  apterous  female.  In  the 
pupae  the  head,  prothorax,  abdomen  and  nectaries  are  of  a  dirty  yellowish  color,  with 
transverse  rows  of  small,  black  or  dusky  spots  on  the  abdomen.  The  wing  pads  are 
black. 

Genus  CERVAPHIS  Van  der  Goot. 

Plate  VIII,  G. 

1916.     Ccrvaphis  Van  dor  Goot,  Zur  Kenntniss  der  Blattliiuse  Java's,  p.  148. 

Characters. — Body  armed  with  a  series  of  long  toothed  projections;  antennas  of 
apterous  form  five-segmented,  of  the  alate  form  six -segmented  with  somewhat  oval 
scnsoria.  Fore  wings  with  the  media  once  branched;  hind  wings  greatly  reduced  in 
size  and  lacking  both  the  media  and  cubitus;  cornicles  elongate,  subcylindric. 
Cauda  and  anal  plate  reduced.  Oviparous  females  often  winged. 

Type  (fixed  by  Van  der  Goot,  191G),  Ccrvaphis  schoutcdcniae  Y.  d.  Goot. 

Subtribe  MACROSIPHINA. 

The  genera  of  the  subtribe  Macrosiphina  may  be  separated  at  once 
from  those  of  the  Aphidina  in  that  they  have  developed  large  antennal 
tubercles.  These  may  assume  various  shapes  by  which  the  genera 
often  may  be  separated.  Considerable  variation  is  met  with  also  in 
the  cornicles  and  cauda,  although  as  a  rule  the  cornicles  are  very  well 
developed.  They  may  be  either  cylindrical  or  swollen;  In  one 
genus,  Hyalopteroides  Theo.,  the  cornicles  are  very  short,  suggesting 
some  of  the  genera  of  the  Aphidina.  The  cauda  is  as  a  rule  rather 
long.  The  wings  are  in  nearly  every  case  normal  in  venation.  The 
different  genera  may  be  separated  by  the  following  key: 

KEY  TO  THE  GENERA  OF  THE  MACROSIPHINA. * 

I .     Cornicles  swollen 2. 

Cornicles  cylindrical  or  tapering,  scarcely  swollen 0. 


1  Since  this  paper  was  set  up  Takahashi  (Insect  World,  v.  23,  p.  439)  has  erected  the  following  genus, 
which  will  fall  in  this  subtribe. 

Genus  AKKAIA  Takahashi. 

Characters. — Cornicles  swollen;  frontal  tubercles  and  Grst  antenn:vl  segment  with  prominent  projec- 
tions.   Anteuuie  of  flvc  segments.    Cauda  somewhat  knobbed,  anal  pluto  lirge  and  projecting. 

Type  (monolypica.}),Akkaia  polyginc,  T.iUaliashi. 


54  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

2.  Cornicles  cylindrical  at  base  and  extremity;  abruptly  swollen  in  middle. . 

Rhopalosiphoninus. 
Cornicles  with  the  swelling  gradual 3. 

3.  Head  with  a  large  central  process  on  vertex Francoa. 

Head  without  this ... .         4. 

4.  Antennal  tubercles  large  and  diverging 5. 

Antennal  tubercles  converging;  head  and  basal  antennal  segments  with 

very  prominent  capitate  hairs Capitophorus. 

5.  Cornicles  much  longer  than  cauda  which  is  somewhat  tapering.  .Amphorophora. 
Cornicles  about  the  length  of  cauda  which  is  usually  constricted  near  its 

base Megoura. 

6.  Cornicles  A-ery  small,  much  smaller  than  the  long,  broad  cauda.  .Ilyalopteroides. 
Cornicles  as  long  as  or  longer  than  the  cauda 7. 

7.  Head  with  prominent,  elongate  projections  to  the  antennal  tubercles,  par- 

ticularly evident  in  the  apterous  form. Phorodon. 

Head  without  these 8. 

8.  Tubercles  strongly  converging Myzus. 

Tubercles  distinctly  diverging 9. 

9.  Cornicles  thick,  about  as  long  as  the  cauda  which  is  large  and  somewhat 

constricted  near  base Macrosiphonella. 

Cornicles  very  long,  rather  slender,  subcylindric,  somewhat  tapering 10. 

10.  Cauda  elongate,  constricted  near  base Macrosiphum . 

Cauda  moderate  or  elongate,  not  constricted  near  base 11 . 

11.  First  antennal  segment  and  abdominal  segments  with  long  fingerlike  tuber- 

cles in  the  apterous  form Acanthaphis. 

Without  these Illinoia. 

Genus  ACANTHAPHIS  Matsumura. 

Plate  VII,  R-U. 
1918.    Acanthaphis  Matsumura,  Trans.  Sapporo  Nat.  Hist.  Soc.,  v.  7,  pt.  1,  p.  15.  . 

The  genus  Acanthaphis  Mats,  is  somewhat  related  to  Phorodon. 

Characters. — Head  with  prominent  diverging  antennal  tubercles.  Antennae  of  six 
segments,  first  segment  with  a  long  fingerlike  projection  in  the  apterous  form.  Corni- 
cles long,  slender,  and  cylindrical.  Cauda  elongate,  conical.  Dorsum  of  abdomen 
with  long  fingerlike  tubercles.  Body  with  capitate  hairs. 

Type  (fixed  by  Matsumura,  1918),  Acanthaphis  rubi  Mats. 

Genus  AMPHOROPHORA  Buckton. 
Plate  IV,  A,  B. 

1876.  Amphorophora  Buckton,  British  Aphides,  v.  1,  p.  187. 

1886.  Macrosiphum  Oestlund,  Minn.  Geol.  Survey  Kept.  14,  p.  27. 

1900.  Macrosiphum  Del  Ouercio,  Nuove  Rel.  Staz.  Firenze,  ser.  1,  no.  2,  p.  159. 

1901.  Nectarosiphon  Schouteden,  Ann.  Ent.  Soc.  Belg.,  v.  45,  p.  112. 
1913.  Eunectarosiphon  Del  Guercio,  Redia,  v.  9,  p.  188. 

1913.    Rhopalosiphum  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  146. 

Buckton  erected  his  genus  Amphorophora  for  his  amputtata,  which 
he  had  secured  in  the  apterous  form  only.  Oestlund  gave  the  generic 
name  Macrosiphum  to  a  species  he  described  as  rubicola.  In  his  sec- 
ond paper  Oestlund  describes  a  species  under  the  name  amputtata 
Buckton  and  says:1  "The  length  of  the  antennae,  together  with  the 
distinct  frontal  tubercles,  may  justify  our  exception  of  Ampho- 
rophora as  a  good  genus."  In  speaking  of  his  Macrosiphum 

i  Bui.  4,  Geol.  and  Nat.  Hist.  Surv.  Minn.,  p.  77, 1887. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  55 

Oestlund  says  it  may  be  too  close  to  Rhopalosiphum  and  compares  it 
with  nymphaeae.  But  had  he  compared  it  with  ampuUata  which  he 
also  included  in  Rhopalosiphum  he  would  probably  have  found  it 
quite  similar,  and  it  is  the  writer's  belief  that  Oestlund's  species  is  in 
reality  Amphorophora. 

Finding  that  Macrosiphum  was  preoccupied  Schouteden  gave  the 
new  name  Nectarosiphon  to  rubicola  and  Nectarosiphon,  therefore, 
will  become  a  synonym.  In  1913,  Del  Guercio  erected  the  genus 
Eunectarosiphon  with  rubi  Kalt.  as  type.  There  appears  not  to  be 
sufficient  difference  here,  however.  The  insects  are  in  their  main 
points  the  same  and  Eunectarosiphon  is  placed,  consequently,  as  a 
synonym.  Van  der  Goot  in  the  same  year  used  ampuUata  as  the 
type  of  Rhopalosiphum. 

In  1900  Del  Guercio  used  the  name  Macrpsiphum  for  a  genus 
including  three  species:  Convolvuli  Kalt.,  viciae  Kalt.,  and  rubi  Kalt. 
He  has  since  used  both  viciae  and  rubi  as  the  types  of  other  genera. 
This  leaves  only  convolvuli  in  his  Macrosiphum,  and  this  species  to  all 
appearances  is  an  Amphorophora.  His  Macrosiphum,  therefore,  is 
listed  here  under  Amphorophora. 

Characters. — Head  with  prominent  and  slightly  diverging  aiitennal  tubercles. 
Antennae  of  six  segments,  armed  with  circular  sensoria.  Wing  venation  normal. 
Cornicles  long  and  somewhat  swollen  in  the  middle.  Cauda  elongate  but  much  dftorter 
than  the  cornicles. 

Type  (monotypical),  Amphorophora  ampuUata  Buckton . 

Genus  CAPITOPHORUS  Van  der  Goot. 

Plate  VIII,  A-C. 
1913.     Capitophorus  Van  der  (loot,  Tijd.  voor  Ent.,  v.  56,  p.  84. 

Iii  the  genus  Capitophorus  Van  der  Goot  certain  species  suggest 
Rhopalosiphum  in  general  characters,  whereas  others  more  nearly 
approach  Myzus  in  their  main  characters.  The  capitate  spines 
appear  to  be  the  best  means  of  determining  the  genus. 

Characters. — Head  with  aiitennal  tubercles  which  are  not  markedly  prominent, 
these  each  with  one  or  more  prominent  knobbed  spines.  Vertex  with  a  central  pro- 
jection on  which  similar  spines  are  located.  Antennae  of  six  segments,  armed  with 
subcircular  sensoria,  the  first  segment  with  a  projecting  process  on  which  one  or  more 
knobbed  spines  are  located.  Wing  venation  normal;  cornicles  long  and  slender, 
slightly  constricted  in  the  middle  and  somewhat  enlarged  toward  the  distal  extremity. 
Cauda  rather  short  and  conical. 

Type  (fixed  by  Van  der  Goot,  1913).  Aphis  carduinm  Walker. 

Genus  FRANCOA  Del  Guercio.i 

1917.     Francoa  Del  Guercio,  Rodin,  v.  12,  p.  201. 

The  genus  Francoa  Del  Guercio  appears  very  close  in  some  respects 
to  Capitophorus  but  on  account  of  the  peculiar  frontal  tubercle  and 
the  structure  of  the  first  aiitennal  segment  it  is  held  to  be  distinct. 

1  It  seems  doubtful  if  this  genus  is  distinct  from  Capitophorus.  We  have  been  unable  to  make  a  careful 
study  of  the  type. 


56  BULLETIN   826,   U.    S.    DEPARTMENT   OF   AGRICULTURE. 

Characters. — Head  with  antenna!  tubercles  present.  Vertex  with  a  prominent  rec- 
tangular process,  both  the  antennal  tubercles  and  the  frontal  process  armed  with 
knobbed  spines.  Antennae  of  six  segments,  the  first  segment  lacking  the  process  and 
capitate  hairs  of  Capitophorus.  Cornicles  rather  slender  and  swollen  near  the  distal 
extremity.  Cauda  elongate,  somewhat  conical. 

Type  (monotypical),  Francoa  elegans  Del  Guercio. 

Genus  HYALOPTEROIDES  Theobald. 

1916.    Hyaloptefoides  Theobald,  The  Entomologist,  v.  49,  p.  51. 

The  genus  Hyalopteroides  was  erected  by  Theobald  for  his  species 
paUida  found  in  the  nest  of  Lasius  niger,  Porlock  Weir,  Somerset. 
It  bears  a  striking  resemblance  to  Pergandeidia  but  there  are  no 
prominent  antennal  tubercles  in  that  genus.  However,  there  are 
slight  swellings  suggestive  of  those  figured  by  Theobald.  The  writer 
has  never  seen  specimens  of  pallida  and  therefore  is  unable  to  give  a 
personal  opinion.  Theobald  says,  "Head  with  marked  frontal 
tubercles."  This  would  place  the  genus  as  not  closely  related  and 
pending  a  study  of  specimens  it  may  be  left  thus. 

Characters. — Head  with  prominent  antennal  tubercles.  Antennae  of  six  segments 
and  armed  with  subcircular  sensoria.  Cornicles  subcylindric,  short,  much  shorter 
than  cauda.  Cauda  long  and  conical. 

Type  (monotypical),  Hyalopteroides  pallida  Theo. 

Genus  ILLINOIA  Wilson. 
Plate  VIII,  H-J. 

1910.  niinoia  Wilson,  Ann.  Ent.  Soc.  Am.,  v.  3,  p.  318. 

1914.     Metopeurum  Mordwilko,  Faune  de  la  Russie,  Hemiptera, ,  v.  1,  p.  56,  67. 
1914.    Acyrthosiphon  Mordwilko,  Faune  de  la  Russie,  Hemiptera,  v.  1,  p.  55,  62. 

This  genus  is  closely  related  to  Macrosiphum  Pass,  from  which  it 
may  be  distinguished  by  the  nature  of  the  cauda. 

Characters. — Head  with  prominent  diverging  frontal  tubercles.  Antennae  of  six 
segments  armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice  branched, 
hind  wings  with  both  media  and  cubitus  present.  Cornicles  cylindrical,  sometimes 
slightly  larger  toward  the  middle  which  appearance  is  accentuated  by  a  constriction 
often  present  near  the  distal  extremity.  Cauda  conical,  not  as  in  Macrosiphum  with 
a  constriction  near  its  base.  Males  usually  winged,  oviparous  females  apterous. 

Type  (fixed  by  Wilson,  1910),  Siphonophora  liriodendri  Mon. 

Genus  MACROSIPHONIELLA  Del  Guercio. 

Plate  VIII,  R-T. 

1911.  Macrosiphoniclla  Del  Guercio,  Redia,  v.  7,  p.  331. 

1913.  Macrosiphum  Van  der  Goot,  Tijd.  voor  Ent.,  v.  56,  p.  145. 

1914.  Diclcysmura  Mordwilko,  Faune  Russ.  Aphidodea,  p.  56. 

The  genus  Macrosiphoniella  was  erected  by  Del  Guercio  with  atrum 
Ferr.  as  type.  In  1913  Van  der  Goot  used  Macrosiphum  of  Passerini, 
indicating  miUefolii  Fab.  as  type.  This  species  can  not  be  made  the 
type  of  Macrosiphum  Pass.,  and  since  it  is  essentially  like  atrum,  V. 
d.  Goot's  Mascrosiphum  must  become  a  synonym  of  Macrosipho- 
niella. In  1914,  Mordwilko  used  the  generic  name  Dielcysmura  for 
miUefolii  and  figured  the  species.  This  name  then  proves  also  to  be 
a  synonym. 


GENERIC    CLASSIFICATION    OF   APHIDIDAE.  57 

Characters. — Head  with  prominent  diverging  antenna!  tubercles.  Antennae  of  six 
segments  armed  with  subcircular  prominent  sensoria.  Fore  wings  with  the  media 
twice  branched,  hind  wings  with  both  media  and  cubitus  present.  Cornicles  short, 
about  the  length  of  the  cauda  and  rather  thick,  usually  with  conspicuous  polygonal 
markings;  cauda  large,  slightly  constricted  near  the  base. 

Type  (fixed  by  Del  Guercio,  1911),  Aphis  atrum  Ferr. 

Ge-us  MACROSIPHUM  Passerini. 
Plate  VIII,  U-W. 

1860.  MacrosipJium  Passcrini,  Gli  Afldi,  p.  27. 

1855.  Siphonophora  Koch,  Die  Pflanzenlaiise  Aphiden,  p.  150. 

1887.  Nectarofiiura  Ocstlund,  Minn.  Geol.  Survey  Bui.  4,  p.  78. 

1913.  IfacrosipJion  Del  Guercio,  Redia,  v.  9,  p.  1S8. 

In  1855  Koch  used  the  generic  name  Siphonophora  for  this  genus 
for  which  Passerini  substituted  Macrosiphum.  Not  aware  of  this 
Oestlund,  seeing  that  Siphonophora  was  preoccupied,  substituted 
Ncctarophora  and  strangely  enough  used  Macrosiphum  for  another 
genus  in  a  different  sense.  Del  Guercio  uses  Macrosiphon  in  this 
sense. 

Characters. — Head  with  prominent  diverging  frontal  tubercles.  Antennte  of  six 
segments,  armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice  branched ; 
hind  wings  with  both  media  and  cubitus  present.  Cornicles  long,  subcylindrical, 
mostly  somewhat  tapering;  cauda  long,  somewhat  constricted  about  the  middle. 
Sexes  with  the  male  usually  alate  and  the  oviparous  form  apterous. 

Type  (fixed  by  Passerini,  1860),  Aphis  rosae  Linn. 

Genus  MEGOURA  Buckton. 

1876.     Megoura  Buckton,  British  Aphides,  v.  1,  p.  188. 
1913.    Drcpaniilli  Del  Guercio,  Rcdia,  v.  9,  p.  188. 

The  genus  Megoura  Buckton  is  similar  to  Amphorophora  in  that 
prominent  antennal  tubercles  are  present.  It  differs,  however,  in 
that  the  cornicles  are  short,  about  equal  in  length  to  the  cauda. 
Buckton  erected  the  genus  for  his  viciae  which  Schoutedeii  con- 
sidered the  same  as  viciae  Kalteiibach.  It  certainly  is  very  similar 
in  every  respect. 

Viciae  Kalt.  was  set  as  the  type  of  his  genus  Drepaiiiella  by  Del 
Guercio,  therefore  Drepaiiiella  will  become  a  synonym  of  Megoura. 

Characters. — Head  with  distinct  antennal  tubercles.  Antenme  of  six  segments, 
armed  with  subcircular  sensoria.  Wing  venation  normal;  cornicles  moderately  long 
and  swollen  in  the  middle.  Cauda  about  the  same  length  as  the  cornicles. 

Type  (rnonotypical),  Megoura  viciae  Buckton. 

Genus  MYZUS  Passerini. 
Plate  VIII,  L,  M,  X-Z. 

1SCO.  Myzu*  Passerini,  Oli  Afldi,  p.  27. 

I860.  Rhopalosiphum  Fasserini,  Gli  Afidi,  p.  27. 

1913.  ITyzoides  Van  der  Goot,  Tijd.  voor  Knt.,  v.  5C>,  p.  SI. 

1913.  Ovalus  Van  dor  Goot,  Tijd.  voor  Ent..  v.  56,  p.  SI. 

10H.  ^^y^odcs  Mordwilko,  Faune  Russ.    Aphidoidea,  p.  52. 

191 1.  AulacorHium  Mordwilko,  Faunc  Russ.  Aphidoidea,  p.  5S. 

1910.  Neomyzus  Van  dcr  Goot,  Zur  Kennt.  der  Blattliiuse  Java's,  p.  50. 

1918.  Myzopsis  Matsumura,  Trans.  Sapporo  Nat.  I  list.  Poo.,  v.  7,  pt.  1,  p.  19. 


58  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

The  genus  Myzus  was  erected  with  Aphis  cerasi  Fab.  as  type. 
In  the  same  year  Passerini  used  ApJiis  persicae  Sulz.  as  type  of 
Rhopalosiphum  Koch.  Aphis  nympJiaeaelj.,  however,  had  been  set 
as  the  type  of  that  genus  in  1856.  Persicae  seems  to  be  very  closely 
related  to  cerasi  and  Rhopalosiphum  (Koch)  Pass.,  therefore,  is 
placed  as  a  synonym  of  Myzus. 

In  1913  Van  der  Goot  placed  certain  other  species  hi  Myzus  and 
the  type  of  the  genus  he  made  the  type  of  his  Myzoides.  This  will 
evidently  then  become  a  synonym,  as  it  has  the  same  type. 

In  1916  Van  der  Goot  erected  the  genus  Neomyzus  with  circum- 
Jlexum  Buckt.  as  type.  While  the  frontal  tubercles  of  this  species  do 
not  converge  quite  to  the  same  extent  as  those  of  cerasi,  they  are 
quite  similar  and  the  cauda  is  somewhat  conical  as  in  that  species. 
Neonrvzus,  therefore,  is  believed  to  be  a  synonym.  Mordwilko  used 
the  name  Myzodes  with  tabaci  Mord.  as  type.  This  species  he  does 
not  fully  describe,  but  gives  a  figure  and  describes  the  characters  of 
the  genus.  From  the  information  given  it  appears  to  be  a  synonym 
of  Myzus.  The  writer  also  believes  that  the  genus  Ovatus  V.  d.  Goot 
is  a  synonym  of  Myzus.  This  genus  was  erected  with  mespili 
V.  d.  Goot  as  type.  The  genus  Aulacorthum  Mord.  was  erected 
with  pelargonii  as  type.  A  study  of  this  species  shows  the  antennal 
tubercles  very  similar  to  those  of  cerasi.  The  cornicles,  too,  are  quite 
similar,  although  the  cauda  is  a  little  more  Aphis-like.  The  writer 
believes  this  genus  is  a  synonym. 

Characters. — Head  with  distinct  antennal  tubercles  present  which,  particularly  in 
the  apterous  form,  project  inward  and  are  strongly  gibbous.  Antennae  of  six  seg- 
ments, the  first  segment  gibbous  like  the  antennal  tubercles.  Wing  venation  normal. 
Cornicles  rather  long  and  subcylindrical.  Cauda  somewhat  short  and  conical,  con- 
stricted very  slightly,  if  at  all. 

Type  (fixed  by  Passerini,  1860),  Aphis  cerasi  Fab. 

Genus  PHORODON  Passerini. 

Plate  VIII,  N-Q. 

1800.    Pharodon  Passerini,  Gli  Afidi,  p.  27, 

Characters. — Head  in  the  alate  form  with  distinct  antennal  tubercles  which  project 
somewhat  inward,  first  antennal  segment  gibbous.  In  the  apterous  form  the  antennal 
tubercles  possess  very  prominent  projections  which  extend  forward  in  front  of  the 
head.  First  antennal  segment  with  a  projecting  process.  Fore  wings  with  media 
twice  branched;  hind  wings  with  both  media  and  cubitus  present.  Cornicles  cylin- 
drical, in  the  apterous  form  somewhat  curved;  cauda  rather  acutely  conical,  not  as 
long  as  the  cornicles. 

Males  as  a  rule  winged  and  oviparous  females  apterous. 

Type  (fixed  by  Passerini,  1860),  Aphis  humuli  Schrk. 

Genus  RHOPALOSEPHONINUS,  n.  gen. 
Plate  IV,  D-F. 

The  genus  Rhopalosiphoninus  is  erected  for  latysiphon  Davidson, 
a  species  with  very  peculiar  cornicles.  It  appears  to  be  somewhat 
related  to  Amphorophora. 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  59 

Characters. — Head  with  prominent  antennal  tubercles  which  project  inward  and  are 
armed  with  prominent  spines.  Anteniue  of  six  segments  armed  with  subcircular 
Bensoria  and  in  the  first  segment  with  spines  similar  to  those  of  the  antennal  tubercles. 
Wing  venation  normal.  Cornicles  narrow  and  cylindrical  at  the  base,  then  abruptly 
and  prominently  swollen,  returning  again  abruptly  to  the  normal  size  near  the  tip. 
Cauda  rather  short  and  conical. 

Type,  Amphorophora  latysiphon  Davidson. 

Subtribe  PENTALONINA. 

The  subtribe  Pentalonina  is  one  of  the  Aphidini  in  which 
specialization  in  the  whig  venation  has  taken  place  in  a  peculiar 
manner.  The  radial  sector  has  in  one  genus  extended  downward 
and  coalesced  with  the  upper  branch  of  the  media.  In  the  genus 
Idiopterus  the  two  have  not  become  entirely  fused,  though  in  some 
specimens  they  have  almost  done  so.  In  Pentalonia,  however,  the 
veins  have  become  permanently  united,  and  a  very  peculiar-looking 
venation  is  the  result.  A  closed  cell  is  formed  by  the  radial  sector 
when  it  meets  the  upper  branch  of  the  media,  and  when  it  leaves 
this  again  it  gives  a  three-branched  appearance  to  the  upper  branch 
of  the  media.  The  explanation  of  this  peculiar  venation  is,  however, 
easily  understood  by  comparison  with  the  venation  of  Idiopterus. 
In  some  of  the  genera  the  hind  wings  are  greatly  reduced,  so  that  some- 
times only  one  vein  remains,  while  in  other  genera  this  reduction  has 
not  taken  place.  Most  of  the  wing  veins  are  clouded  with  brownish 
borders. 

The  insects  feed  usually  upon  ferns  or  tropical  plants. 

KEY  TO  THE  GENERA  OF  THE  PENTALONINA. 

1 .  Hind  wings  much  reduced  in  size,  lacking  at  least  the  cubitus 2. 

Hind  wings  nearly  normal  in  size  and  with  both  media  and  cubitus  present.         3. 

2.  Radial  sector  of  fore  wing  fused  with  the  upper  branch  of  the  media,  forming 

a  closed  cell Pentalonia. 

Radial  sector  of  fore  wings  not  so  fused,  but  normal Microparsus. 

3.  Cornicles  cylindrical Idiopterus. 

Cornicles  somewhat  swollen  near  their  distal  extremities 4. 

4.  Media  of  fore  wings  twice  branched Fullawayella. 

Media  of  fore  wings  once  branched Keotoxoptera. 

Genus  FULLAWAYELLA  Del  Guerclo. 

1011.     Fullau-ayclla  Del  Oucrcio,  Redia,  v.  7,  p.  462. 

191t>.     Micrnmyzus,  Van  dor  Goot,  Zur  Kenntniss  dor  Blattlaiisc  Java's,  p.  53. 

This  genus  is  very  suggestive  of  Amphorophora  in  certain  ways 
but  no  doubt  is  related  here.  Van  der  Goot's  genus  was  erected 
with  nigrum  V.  d.  Goot  as  type  but  this  species  differs  very  little 
from  Jcirkdldyi.  . 

Characters. — Antenna;  on  prominent,  converging,  imbricated  antennal  tubercles,  of 
six  segments  and  armed  with  subcircular  sensoria.  Fore  wings  with  the  media  twice 
branched  and  with  the  radical  sector  deeply  curved  toward  the  upper  branch  of  the 


60  BULLETIN"   826,   TJ.    S.    DEPARTMENT   OF   AGRICULTURE. 

media;  hind  wings  with  both  media  and  cubitus  present.     Cornicles  somewhat  swollen 
near  their  distal  extremities.     Cauda  elongate  and  constricted  near  the  base. 

Type  (mono typical),  Macrosiphum  Tcirkaldyi  Fullaway. 

Genus  IDIOPTERUS  Davis. 
Plato  VIII,  EE-HH. 

1900.    Idiopterus  Davis,  Ann.  Ent.  Soc.  Amer.,  v.  2,  p.  193. 

Idiopterus,  a  less  specialized  genus  than  Pentalonia,  is  worthy  of 
special  note,  as  it  gives  a  key  to  the  peculiar  venation  of  the  latter  genus. 
The  coalescing  of  the  radial  sector  and  the  media  is  here  plainly 
visible  and  in  some  specimens  a  triangular  closed  cell  is  formed, 
although  in  most  examples  the  two  veins  can  be  traced  distinctly. 

Characters.  —Head  with  prominent  antennal  tubercles  which  project  slightly  in- 
wards, and  are  gibbous.  Antennae  of  six  segments,  armed  with  subcircular  sensoria, 
the  first  segment  gibbous  like  the  antennal  tubercles.  Corniclea  subcylindric,  rather 
slender,  cauda  somewhat  elongate,  conical.  Fore  wings  with  the  radial  sector  extend- 
ing abruptly  downward  from  the  stigma  and  paralleling  the  upper  branch  of  the 
media  with  which  in  some  specimens  it  appears  to  be  almost  united ;  hind  wings  with 
both  media  and  cubitus  present. 

Type  (mono typical),  Idiopterus  nephrolepidis  Davis. 

Genus  MICROPARSUS  Patch. 

Plate  VIII,  AA-DD. 
1909.     Microparsus  Patch,  Ent.  News,  v.  20,  p.  337. 

Microparsus  is  at  once  distinguished  from  the  other  genera  related 
to  it  by  the  peculiar  venation  and  the  reduction  of  the  hind  wing. 

Characters. — Head  with  distinct  antennal  tubercles  present.  Antenna;  of  six  seg- 
ments, armed  with  subcircular  sensoria.  Fore  wings  with  the  media  once  branched; 
hind  wings  much  reduced  in  size  and  lacking  both  the  media  and  cubitus.  Cornicle's 
subcylindric.  Cauda  rather  long  and  tapering,  almost  equal  in  length  to  the  cornicles. 

Type  (monotypical),  Microparsus  variabilis  Patch. 

Genus  NEOTOXOPTERA  Theobald.i 

1915.     Ncotoxopicra  Theobald,  Bui.  Ent.  Res.,  v.  6,  p.  131. 

This  genus  is  closely  related  to  Fullawayella,  from  which  it  can  be 
separated  by  the  venation.  There  has  been  some  doubt  whether  or 
not  this  is  a  good  genus,  for  the  name  will  not  hold  if  the  type  is  found 
to  correspond  with  Pergande's  violae,  which  it  resembles.  In  that 
case  Neotoxoptera  would  become  a  synonym  of  Fullawayella,  for 
Pergande's  species  is  undoubtedly  a  Fullawayella. 

Characters. — Head  with  prominent  antennal  tubercles.  Antenme  of  six  segments, 
armed  with  subcircular  sensoria.  Fore  wings  with  the  media  once  branched.  Corni- 
cles swollen,  elongate,  somewhat  conical. 

Type  (monotypical),  Neotoxoptera  violae  Theo. 

i  After  this  paper  v.-asin  lype  the  writer  (Bui.  Ent.  Res.,  v.  10,  p.  45)  showed  that  violae  Theo.  is  a 
synonym  of  violae  1'erg. 


GENERIC   CLASSIFICATION   OF  APHIDIDAE.  61 

Genus  PENTALONIA  CoquereL 

Plato  VIII,  II-MM. 
1859.    Pentalonia  Coquerel,  Ann.  Ent.  Soc.  France,  Ser.  3,  v.  7,  p.  259. 

The  genus  Pentalonia  Coquerel  is  a  very  peculiar  one  and  possesses 
a  venation  unlike  that  of  any  other  in  the  Aphididae.  It  is,  however, 
only  a  little  further  development  of  the  condition  met  with  in 
Idiopterus,  which  is  the  less  specialized  of  the  two  genera. 

Characters. — Head  with  prominent  antennal  tubercles  which  are,  more  especially 
in  the  apterous  form,  projected  inward,  gibbous  and  somewhat  Myzus-like  in  appear- 
ance. Antennae  of  six  segments,  armed  with  subcircular  sensoria,  the  first  segment 
gibbous  like  the  antennal  tubercles.  Cornicles  somewhat  constricted  near  their 
middle,  then  again  somewhat  swollen  near  their  distal  extremity.  Cauda  rather  small 
but  elongate,  subconical,  slightly  constricted  about  the  middle.  Fore  wings  with  the 
radial  sector  extending  abruptly  downward  and  meeting  the  upper  branch  of  the  media 
with  which  it  fuses  but  is  diverted  again  toward  its  natural  course  near  the  tip  of  the 
wing.  A  closed  cell  is  thus  formed  by  the  radial  sector  and  the  media  but  at  the 
margin  of  the  wing  there  are  the  same  veins  as  in  the  Aphidini  (Plate  VIII,  JJ.) 
Hind  wings  very  much  reduced,  cubitus  absent. 

Type  (monotypical),  Pentalonia  nigronervosa  Cql. 

Subfamily  II,  MINDARINAE. 

It  has  been  the  custom  to  consider  the  genus  Mindarus  as  closely  re- 
lated to  the  Pemphigini,  but  the  writer  is  unable  to  do  this  and  concludes 
that  it  must  represent  a  subfamily  in  itself.  In  some  ways  abietinus 
is  the  most  primitive  living  aphid.  It  is,  in  fact,  the  only  one  which 
has  retained  the  general  wing  structure  which  is  predominant  in  the 
fossil  forms.  It  is  true  that  the  venation  is  more  reduced  than  in 
some  of  the  other  subfamilies,  but  the  type  of  wing  in  regard  to  the 
stigma  formation  is  exactly  like  most  fossil  wings  and  unlike  the 
wings  of  other  living  forms.  Many  of  the  characters  suggest  the 
Eriosomatinae  and  the  genus  is  no  doubt  very  similar  to  the  ancestors 
of  the  insects  in  that  subfamily.  The  antennal  structure  and  general 
form  are  like  those  in  the  Eriosomatinae.  The  sexes,  too,  are  apterous, 
but  though  they  have  developed  the  small  apterous  condition  they  are 
in  many  ways  more  primitive  than  are  the  sexes  of  the  Eriosomatinae. 
The  male  is  small  and  suggests  the  condition  in  those  forms.  The 
peculiar  habit  of  copulation  is  similar,  in  that  the  male  mounts  the 
female  and  may  remain  there  inactive  for  a  very  long  period.  The 
writer  has  observed  a  male  of  Eriosoma  lanigerum  clinging  thus  to  a 
female  for  48  hours.  The  sexes  of  Mindarus,  however,  have  not 
lost  the  beak  and  the  male  feeds  on  the  juices  of  its  host.  In  this 
regard  they  are  more  primitive  than  sexes  in  the  Eriosomatinae. 
The  oviparous  female,  moreover,  develops  her  ovaries  and  produces 
as  high  as  8  or  9  eggs,  in  striking  contrast  with  the  ovipara  in  the 
Eriosomatinae.  It  is  a  much  less  specialized  condition.  In  regard 
to  the  alate  form  the  shape  of  the  cauda  is  quite  different  from  that 
met  with  in  the  Eriosomatinae. 


62  BULLETIN   826,   U.   S.   DEPARTMENT   OF  AGRICULTURE. 

It  seems  to  the  writer  that  the  Mindariiiae  give  a  fair  idea  of  the 
ancestors  of  the  Eriosomatinae  'and  may  even  represent  a  group 
dominant  in  earlier  times  from  which  the  Eriosomatinae  sprang. 

Only  one  genus  is  represented. 

Genus  MINDARUS  Koch. 

Plate  IX,  A-F. 
1857.  Koch,  Die  Pflanzenlause  Aphidcn,  p.  277. 

The  peculiar  genus  Mindarus  was  erected  by  Koch  with  abietinus 
Koch  as  type.  This  species  is  the  only  one  in  the  genus,  although  it 
has  been  redescribed  as  Schizoneura  pinicola  Thos.  and  Schizoneura 
oUiqua  Choi. 

Characters. — Cornicles  present  as  mere  rings.  Large  wax  plates  present.  Alate 
forms  with  six-segmented  antennae  armed  with  oval  sensoria.  Fore  wings  with  the 
media  once  branched;  radial  sector  inserted  mesad  of  the  long  narrow  stigma,  thus 
igiving  a  very  long  stigmal  cell;  bind  wings  with  both  media  and  cubitus  present. 
Cauda  rather  long,  not  rounded,  but  somewhat  conical  or  even  spatulate.  Sexes  email 
and  apterous,  beaks  present  and  feeding  taking  place.  Oviparous  female  with  the 
ovaries  developed  and  laying  as  high  as  9  eggs.  Forms  living  free  upon  the  twigs  of 
conifers  which  become  somewhat  distorted  by  the  feeding  of  the  insects. 

Type  (monotypical),  Mindarus  abietinus  Koch. 

Subfamily  HI,  ERIOSOMATINAE. 

The  subfamily  Eriosomatinae  is  composed  of  insects  which  are 
perhaps  as  specialized  as  any  of  the  Aphididae.  They  show  a  re- 
markable  development  of  the  habit  of  gall  formation  and  in  this 
respect  parallel  the  Hormaphidinae.  The  insects  of  that  subfamily, 
however,  evidently  have  developed  the  habit  independently.  Many 
previous  authors  have  placed  all  of  these  forms  in  the  present  sub- 
family. This,  the  writer  believes,  is  incorrect,  as  shown  by  the  biol- 
ogies of  the  insects.  The  sexual  forms  give  a  true  understanding  of 
the  relationships  and  of  the  genera  which  should  be  included  in  the 
Eriosomatinae.  All  of  the  forms  included  by  the  writer  show  evi- 
dence of  a  common  origin  in  that  the  sexes  have  become  degenerate. 
They  have  become  small  apterous  forms  and  have  lost  the  mouth  parts 
and  the  ability  to  take  food.  That  this  was  not  their  original  condition 
is  clearly  shown  by  the  history  of  the  family  and  also  by  the  fact  that 
the  sexual  forms  of  some  species  have  a  beak  when  born,  but  lose  this 
at  the  first  molting.  Other  species  even  at  the  time  of  birth  are  devoid 
of  all  but  a  rudimentary  trophictubercle.  The  reproductive  system 
of  the  female  has  become  greatly  altered.  As  previously  pointed 
out  by  the  writer,  the  early  development  of  the  reproductive  system 
of  the  sexual  female  corresponds  exactly  to  that  in  the  apterous  forms 
and  to  that  of  the  oviparous  forms  of  the  more  primitive  groups. 

Young  embryos  *  *  *  show  that  the  ovaries  are  at  first  similar  to  those  of  the 
parthenogenetic  form.  There  may  be  distinguished  the  four  chambers  on  each  side 
containing  egg  cells  and  nutritive  cells.  In  later  embryos  most  of  the  egg  tubes  arc  in 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  63 

the  process  of  degeneration  and  only  two  ovaries,  one  on  each  side,  develop.  Of  these 
one  finally  degenerates  and  the  egg  of  the  other  grows  until  it  fills  almost  the  entire 
body  and  the  insect  appears  to  be  little  else  than  egg.1 

It  will  be  seen  at  a  glance  that  such  a  method  of  egg  development 
is  entirely  different  from  that  met  with  in  members  of  the  genera  which 
have  been  heretofore  placed  in  the  subfamily.  The  Hormaphidini  and 
the  Thelaxini,  as  will  be  seen  under  the  discussion  of  those  tribes, 
have  sexual  females  which  develop  normal  ovaries  and  lay  several 
eggs  in  the  same  way  as  do  the  Aphidini,  Lachnini,  and  other  groups. 
It  is  true  that  some  have  developed  gall  formation  and  highly  special- 
ized, wax-secreting  organs,  but  this  is  more  of  a  parallelism  than  a 
close  relationship,  as  is  indicated  by  the  sexual  forms.  The  wax- 
secreting  organs  of  the  Eriosomatinae  vary  considerably  in  structure. 
A  study  of  those  in  the  genus  Eriosoma  has  been  presented  by  the 
writer  (1915).  The  glands  here  are  compound,  each  cell  containing  a 
central  wax  chamber  into  which  the  wax  is  secreted  and  from  which 
it  is  forced  out  as  a  fine  waxen  thread.  In  other  genera  the  wax 
glands  take  on  the  nature  of  plates,  illustrated  in  the  genus 
Prociphilus.  These  glands  are  essentially  the  same  in  general  struc- 
ture as  are  those  in  Eriosoma,  but  the  wax  cells  are  placed  very 
close  together  and  are  so  extremely  elongate  that  their  openings  to  the 
surface  are  very  small.  A  large  number  of  these  gives  the  appearance 
of  a  more  or  less  uniform  plate.  The  structure,  however,  in  the  two 
genera  follows  the  same  lines. 

The  wing  venation  in  this  subfamily  presents  as  great  a  reduction 
as  in  any  of  the  subfamilies  of  the  Aphididae  and  in  this  respect  it  is 
comparable  to  the  Hormaphidinae.  In  the  fore  wings  the  reduction 
is  shown  in  the  media  which  is  never  branched  more  than  once.  Dr. 
Patch  has  pointed  out  the  homologies  of  the  veins  and  has  indicated 
that  in  all  of  these  cases  the  branches  represent  M1+2  and  M3+4. 
In  some  cases,  however,  it  would  appear  as  if  they  were  Mt  and  M4. 
In  other  genera  the  media  is  indicated  as  a  single  vein.  The  radial 
sector  is  in  nearly  every  case  present  and  the  cubitus  and  first  anal 
are  prominent  veins.  The  tracheae  are  figured  for  the  subfamily 
under  the  genus  Eriosoma.  In  the  hind  wings  the  radial  sector  is 
always  present  and  two  oblique  veins  are  nearly  always  found. 
These  are  the  media  and  the  cubitus.  In  several  genera,  however, 
the  cubitus  has  disappeared  and  only  the  media  remains  as  the  one 
transverse  vein  in  the  hind  wings. 

The  cornicles  in  the  genera  of  this  subfamily  are  not  prominently 
developed.  Indeed,  they  are  absent  altogether  in  certain  of  the 
tribes.  In  the  genus  Eriosoma  they  are  chitinized  rings  slightly 
elevated  on  shallow  hairy  cones.  The  opening  of  the  cornicles  is 
closed  by  a  muscle  and  from  the  cornicle  a  narrow  duct  leads  to  a 

1  Baker,  A.  C.    The  woolly  apple  aphi.s.     I'.  S.  Dcpt.  Agr.,  OH.  of  Sec.,  Kept.  101,  p.  43.    1915. 


64  BULLETIN   826,   U.   S.   DEPARTMENT  OF   AGRICULTURE. 

large  wax  reservoir.  The  structure  of  the  cornicles  themselves  in  this 
subfamily  is  essentially  the  same  in  all  genera  where  they  are  present. 
In  a  large  number  of  genera,  however,  the  wax  reservoir  is  absent 
and  in  some  specialized  tribes  the  cornicles  are  likewise  absent.  It  is 
interesting  to  note  that  in  some  genera,  though  absent  in  the  stem 
mother,  they  are  present  in  the  alate  forms. 

The  habit  of  gall  formation  is  not  found  equally  in  all  genera 
and  it  would  seem  that  those  forms  which  have  become  associated 
with  ants  have  not  developed  this  habit  to  the  same  extent  as  have 
some  of  the  other  groups.  However,  it  must  be  borne  in  mind  that  our 
knowledge  concerning  the  species  associated  with  ants  is  very  incom- 
plete, and  the  writer  is  convinced  that  many  of  the  Prociphilini  will 
be  found  during  their  summer  generations  in  this  relation.  Many 
of  the  forms  cause  true  galls  which  are  the  result  of  outgrowths 
of  the  plant  and  which  completely  enclose  the  insects.  Sometimes 
the  stem  mother  lives  in  a  gall  by  herself  while  in  other  cases  the 
following  generations  live  with  her.  The  original  spring  gall  is  usually 
the  result  of  the  activities  of  the  young  stem  mother.  Certain  species 
do  not  produce  true  galls  but  form  pseudogalls  which  are  due  to  the 
rolling  or  crumpling  of  the  leaves  on  which  the  insects  feed.  Other 
species,  again,  especially  during  their  summer  generations,  feed  011  the 
twigs  or  roots  of  plants  and  give  rise  to  excrescences  by  their 
feeding..  It  often  happens  that  species  which  in  their  spring  forms 
are  gall  makers,  attack  plants  in  this  way  in  their  summer  generations. 
Others  live  on  the  roots  of  grasses  during  these  generations  and  do 
not  cause  the  excrescences  produced  by  those  species  feeding  on  trees 
and  woody  shrubs. 

The  association  with  ants  is  highly  developed  by  one  tribe  of  this 
subfamily,  although  all  of  the  other  tribes  are  to  a  degree  tended  by 
these  insects.  The  species  of  the  Fordini  live  exclusively  in  the  nests 
of  ants  or  are  tended  by  them,  and  they  are  cared  for  very  carefully 
in  return  for  the  honeydew  excreted.  Ants  also  attend  species  which 
have  aerial  feeding  habits  and  they  may  be  seen  carrying  the  root 
generations  of  species  of  Eriosoma  from  one  place  to  another  and 
even  distributing  them  about  on  the  trees.  Indeed  the  writer  once 
took  advantage  of  the  presence  of  ants  to  infest  some  apple  seed- 
lings. A  vial  of  apterous  insects  was  emptied  at  the  base  of  each 
tree  and  the  ants  soon  could  be  seen  running  about  carrying  the 
aphids  to  suitable  positions  on  the  trees.  Sometimes,  however,  they 
carried  them  away. 


GENERIC   CLASSIFICATION   OF   APHIDIDAE.  65 

KEY  TO  THE  TRIBES  OF  THE  ERIOSOMATINAE. 

1.  Cornicles  present,  at  least  in  the  alate  forms;  however,  often  mere  rings 4. 

Cornicles  absent 2. 

2.  Forms  living  in  true  galls  or  in  pseudogalls  on  plants 3. 

Forms  living  in  the  nests  of  ants  or  at  least  subterranean,  feeding  on  the 

roots  of  plants;  wax-secreting  areas  present;  antennae  of  alate  forms  rather 
short  and  thick  with  somewhat  oval  sensoria FORDINI. 

3.  Forms  living  in  true  galls  and  without  wax  plates  prominently  developed 

on  the  head  and  thorax  of  alate  form;  wax-secreting  areas  present  but  not 
prominently  developed.  Alate  forms  leaving  the  galls  in  the  late  summer  or 

fall.    Antennae  with  annular  sensoria MELAPHINI. 

Forms  living  in  pseudogalls,  occasionally  in  true  galls.  Antennae  of  alate 
form  rather  long  and  slender,  with  narrow  or  somewhat  oval  or  rounded 
sensoria .  Wax  plates  well  developed  and  present  on  the  head  and  thorax 
of  the  alate  forms  which  leave  the  galls  in  the  spring PROCIPHILINI. 

4.  Forms  living  in  galls,  pseudogalls,  or  free  upon  their  host; -wax  glands  prom- 

inently developed;  antennae  of  alate  forms  armed  with  annular  sensoria 

which  almost  completely  encircle  the  segments ERIOSOMATINI. 

Forms  living  usually  in  true  galls;  wax  glands  present  but  not  strongly  de- 
veloped; antennae  of  alate  forms  armed  with  narrow,  transverse  sensoria, 
somewhat  oval  or  irregular  ones,  or  occasionally  without  sensoria.  .PEJIPHIGINI. 

Tribe  ERIOSOMATINI. 

The  tribe  Eriosomatini  is  composed  of  insects  which  have  more 
or  less  developed  the  habit  of  gall  formation,  which  are  possessed  of 
wax  glands,  and  the  antennae  of  the  alate  forms  of  which  are  armed 
usually  with  annular  sensoria.  Their  typical  host  group  is  that  of 
the  elms. 

Characters. — Forms  living  in  galls,  pseudogalls,  or  free  upon  the  twigs  or  roots  of 
their  host  on  which  they  form  excrescences.  Prominent  wax  glands  present.  Cor- 
nicles distinct;  antennaeof  alate  forms  armed  with  annular  sensoria  which  often  almost 
completely  encircle  the  segments.  Sexual  forms  small,  apterous,  beakless;  oviparous 
females  developing  a  solitary  egg. 

KEY  TO  THE  GENERA  OF  THE  ERIOSOMATIVI. 

1 .  Media  of  the  fore  wings  of  alate  form  once  branched 3. 

Media  of  the  alate  form  simple 2. 

2.  Hind  wings  with  both  media  and  cubitus  present Gobaishia. 

Hind  wings  with  only  the  media  present Tctraneura.1 

3.  Hind  wings  with  both  media  and  cubitus  present Eriosoma. 

Hind  wings  with  only  the  media  present 4. 

•1.     Stem  mother  with  four-segmented  antennae;  antennre  of  alate  form  rather 

short  and  thick Colopha. 

Stein  mother  with  five-segmented  antenna1 ;  antennae  of  alate  form  long  and 
slender Georgia. 

i  There  is  considerable  evidence  for  separating  a  tribe  Tctraneurini  to  include  the  genera  Colopha, 
Tetraneura,  and  Gobaishia. 

141G1.S0— 20— Bull.  82G 5 


66  BULLETIN  826,  U.  S.  DEPARTMENT  OF  AGRICULTURE. 

Genus  COLOPHA  Monell. 

Plate  IX,  G-L. 
1877.     Cotopha  Monell,  Can.  Ent.,  v.  fi,  p.  102. 

The  genus  Colopha  was  erected  for  ulmicola  Fitch.  One  of  the 
principal  characters  whereby  it  may  be  separated  from  Tetraneura 
is  the  once-branched  character  of  the  media.  The  two  genera,  how- 
ever, are  very  closely  related.  The  species  have  the  same  peculiar 
structure  and  the  same  mode  of  life. 

Characters. — Cornicles  slightly  elevated  rings.  Stem  mother  with  four-segmented 
antennse.  Apterous  form  with  five-segmented  or  sometimes  six-segmented  antennse. 
Wax  glands  present.  Alate  form  with  six-segmented  antennae  which  are  armed  with 
annular  sensoria  partly  encircling  the  segments.  Fore  wings  with  the  media  once 
branched,  hind  wings  with  only  the  media  present.  Forms  making  galls  upon  the 
leaves  of  trees  in  which  the  stem  mother  and  her  offspring  live  in  company;  in  summer 
migrating  to  the  roots  o«f  plants. 

Type  (monotypical),  Byrsocrypta  ulmicola  Fitch. 

Genus  ERIOSOMA  Leach. 
Plate  IX,  M-T. 

1818.  Eriosoma  Leach,  Trans.  Hort.  Soc.  London,  v.  3,  p.  60. 

1831.  Myzoxylus  Blot,  Mem.  Soc.  Roy.  Agr.  et  de  Com.  Caen.,  v.  3,  p.  332.  » 

1837.  Schizoneura  Hartig,  Jahresb.  il.  d.  Fortschr.  d.  Forstwiss.  und  forstl.  Naturk.,  v.  1,  p.  645. 

1848.  Mimaphidus  Rondani,  Nuovi  Annali  della  Seienze  Natural!,  ser.  2,  v.  9,  p.  35. 

In  1818  Leach  erected  his  genus  hi  a  footnote  in  connection  with  a 
paper  read  by  Mosley.  The  paper  was  published  in  1818.  In  1819 
Samouelle  published  his  "Useful  Compendium"  and  on  page  232 
characterized  the  genus  Eriosoma  Leach  MSS.  The  printed  copy 
of  the  Transactions  appeared  complete  in  1820.  In  1824  Blot  used 
the  word  Myzoxyle  which  he  corrected  to  Myzoxylus  in  1831.  For 
these  Aphis  lanigera  Hausm.  was  used  as  type. 

In  1837  Hartig  erected  Schizoneura  and  of  this  genus  ulmi  was  made 
type  by  Passerini  in  1860.  This  then  will  become  a  synonym. 
Corni  Fab.  was  for  a  time  considered  the  type  of  this  genus  but  this 
species  was  not  in  the  original  genus.  In  1848  Rondani  used  Mima- 
phidus with  ulmi  Fab.  as  type,  which  according  to  Passerini  is  the 
same  as  lanuginosa  Hartig.  Therefore,  this  genus  will  become  a 
synonym. 

Characters. — Cornicles  distinct  rings  on  somewhat  elevated  tubercles.  Apterous 
form  with  six-segmented  antennae.  Stem  mother  with  five-segmented  antennas. 
Wax  plates  present  in  the  apterous  and  alate  vivipara.  Alate  form  with  six-seg- 
mented antennae  armed  with  annular  sensoria.  Fore  wings  with  the  media  once 
branched,  hind  wings  with  both  media  and  cubitus  present;  cauda  and  anal  plate 
rounded.  Forms  living  in  gall-like  formations  or  causing  excrescences  on  their  hosts. 
Sexual  forms  small,  apterous,  beakless.  Only  one  egg  of  those  of  the  oviparous  female 
develops. 

Type  (monotypical),  Aphis  lanigera  Hausmann. 


GENERIC  CLASSIFICATION  OF  APHIDIDAE.  67 

Genus  GEORGIA  Wilson. 

Plate  IX,  U-Z. 
1911.     Georgia  Wilson,  Can.  Ent.,  v.  43,  p.  64. 

The  genus  Georgia  appears  to  be  related  to  Colopha  Mon.,  and  yet 
many  of  the  characters  are  so  like  those  of  Eriosoma  that  the  insect 
suggests  that  genus  also.  Especially  to  the  species  E.  americanum 
Riley  there  is  a  striking  resemblance.  Prominent  wax  glands  are 
lacking  but  these  are  sometimes  also  lacking  in  the  spring  forms  of 
Eriosoma. 

Characters. — Cornicles  present  and  situated  on  shallow  hairy  cones  as  are  those  of 
Eriosoma.  Stem  mother  with  five-segmented  antennae ;  alate  form  with  six-segmented 
antennye  which  are  armed  with  narrow  sensoria  that  do  not  encircle  the  segment  to 
any  extent.  Fore  wings  with  the  media  once  branched,  hind  wings  with  only  one 
oblique  vein.  Cauda  rounded.  Prominent  wax  pores  such  as  those  present  in 
Eriosoma  lacking,  but  small  wax  areas  present. 

Forms  living  in  pseudogalls  on  plants,  the  alate  individuals  migrating  from  the 
galls  in  the  early  spring. 

Type  (monotypical),  Georgia  ulmi  Wlsn. 

Genus  GOBAISHIA  Matsumura. 
Tlate  X,  A-G;  XI,  V. 

1909.    Byrsocrypta  Tullgren,  Arkiv  for  Zoologi,  Bd.  5,  no.  14,  p.  182. 

1917.    Gobaishia  Matsumura,  Synopsis  of  the  I'emphigidae  of  Japan,  Gifu,  Japan,  p.  75. 

Tullgren  used  the  name  Byrsocrypta  Hal.  as  the  name  of  a  sub- 
genus  with  paUida  as  type,  placing  it  under  Tetraneura.  He  appar- 
ently overlooked  the  fact  that  Westwood  had  set  bursarius  as  the 
type  of  the  genus  Byrsocrypta  as  will  be  found  discussed  under  the 
genus  Pemphigus.  Pallida  is  different  from  the  species  of  Tetraneura, 
in  that  the  cubitus  is  retained  in  the  hind  wing.  If  Colopha  is  retained 
on  account  of  the  branched  nature  of  the  media  in  the  fore  wings  it 
will  be  necessary  to  place  pallida  as  typical  of  a  genus  related  to 
Tetraneura. 

In  1917  Matsumura  erected  the  genus  Gobaishia  with  Gobaishia 
japonica  Mats,  as  type.  This  species  was  stated  to  be  very  similar  to 
Tetraneura  alba  Ratz.  Tetraneura  alba  Ratz  is  the  same  species  as 
Eriosoma  paUida  Haliday  and  the  characters  given  for  the  genus  are, 
therefore,  similar  to  Tullgren's  conception  of  Byrsocrypta.  The 
figures  drawn  are  from  specimens  of  paUida  as  no  japonica  was  avail- 
able to  the  writer  for  study. 

Characters. — Cornicles  present,  stem  mother  with  four-segmented  antenna?,  alate 
forms  with  six-segmented  antennae  which  are  armed  with  annular  sensoria.  Fore  wings 
with  the  media  usually  simple;  hind  wings  with  both  cubitus  and  media  present. 

Type  (fixed  by  Matsumura,  1917),  Gobaishia  japonica  Mats. 


68  BULLETIN  826,  U.  S.  DEPARTMENT  OF  AGRICULTURE. 

Genus  TETRANEURA  Hartlg. 

Plate  X,  H-M. 
1841.     Tetraneun  Hartig,  Germar's  Zeitschrift  fur  die  Entomologie,  v.  3,  p.  366. 

In  1841  Hartig  erected  the  genus  Tetraneura  under  which  he  gave 
Tetraneura  ulmi  Lin.  ?  questioned  thus  and  described.  He  also 
listed  T.  rugicornis  Hartig.  One  of  these  species  was  questioned 
and  the  other  merely  listed.  Ulmi  L.,  however,  was  questioned  only 
in  the  sense  of  the  determination,  and  a  good  description  was  given 
so  that  it  is  known  what  insect  Hartig  had. 

In  1843  Kaltenbach  gave  a  description  of  the  genus  Tetraneura 
crediting  it  to  Hartig  and  described  thereunder  one  species,  Aphis  ulmi 
De  Geer.  ApJiis  ulmi  De  Geer  (1773)  is  the  same  species  as  Aphis 
ulmi  Geoffrey  (1764)  but  this  name  can  not  be  used,  since  Linnaeus 
used  Aphis  ulmi  for  a  different  insect.  This  is  the  same  insect  de- 
scribed by  Hartig  as  T.  ulmi  L.  ?  and  it  is  evident  that  it  requires  a 
new  name,  to  which  ulmifoliae  is  given. 

Characters. — Cornicles  very  slightly  elevated  rings,  not  at  all  prominent.  Stem 
mother  with  four-segmented  antennae;  apterous  form  with  five-segmented  antennae. 
Wax  glands  present.  Alate  form  with  six-segmented  antennae  which  are  armed  with 
narrow  annular  sensoria  almost  completely  encircling  the  segment.  Fore  wings  with 
the  media  simple;  hind  wings  with  only  the  media  present. 

Forms  living  in  galls  and  migrating  in  spring  to  other  plants.  Sexes  small,  apterous 
and  beakless.  Oviparous  female  developing  only  one  egg. 

Type,  Tetraneura  ulmifolix  Baker  (Aphis  ulmi  L.  of  ITartig). 

Tribe  PEMPHIGINI. 

The  tribe  Pemphigini  is  composed  of  forms  which  are  highly  spe- 
cialized and  most  of  which  have  developed  the  habit  of  true  gall 
formation.  The  secretion  of  wax  also  occurs  but  wax  secreting  plates 
are  not  developed  to  the  extent  met  with  in  some  of  the  other  tribes 
of  the  subfamily.  Alternation  of  hosts  is  found  to  occur,  migrants 
leaving  the  galls  in  early  spring  or  summer  and  returning  in  autumn. 
In  some  species,  however,  the  insects  do  not  leave  the  galls  until  the 
mothers  of  the  sexual  forms  -are  produced.  Distinct  cornicles  are 
present  and  by  this  character  forms  in  some  of  the  other  tribes  which 
are  suggestive  of  the  Pemphigini  may  be  distinguished.  The  typical 
host  group  is  Populus  and  the  galls  are  normally  spring  galls. 

Characters. — Forms  usually  inhabiting  true  galls  and  often  migrating  to  other  plants 
during  the  summer.  Antennae  of  six  segments  in  the  alate  form  and  in  nearly  all 
genera  armed  with  linear,  oval,  or  somewhat  irregularly  shaped  sensoria.  Small  wax- 
secreting  areas  present.  Sexual  forms  small,  apterous,  and  'beakless,  the  oviparous 
female  developing  only  one  egg. 

Six  genera  may  be  included  in  the  tribe  and  these  genera  may  be 
separated  by  the  following  key : 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  69 

KEY  TO  THE  GENERA  OF  THE  PEMPHIGINI. 

1.  Unguis  of  segment  VI  of  alate  form  distinctly  long  and  Aphis-like.  .Mordwilkoja. 
Unguis  short  and  knob-like 2. 

2.  Media  once  branched 3. 

Media  simple 4. 

3.  Antennae  of  alate  form  usually  without  secondary  sensoria.     Winga  flat 

in  repose Phloeomyzw. 

Antennae  of  alate  form  with  secondary  sensoria.     Wings  not  flat  in  repose 

Pachypappella . 

4.  Both  media  and  cubitus  present  in  hind  wing 5. 

One  oblique  vein  only  in  hind  wing Dryopeia. 

5.  Antennae  of  alate  form  without  secondary  sensoria.     Wings  flat  in  repose 

Rhizoctonus. 

Antennae  of  alate  form  with  secondary  sensoria.     Wings  not  flat  in  repose. . .         G. 

6.  Antennae  of  alate  form  rather  short  and  thick.     Stem  mother  with  four-seg- 

mented antennae Pemphigus. 

Antennae  of  alate  form  rather  long  and  slender.     Stem  mother  with  five-seg- 
mented antennae Cornaphis. 

Genus  CORNAPHIS  Gillette. 

Plate  X,  N-T. 
1913.     Cornaphis  C.illette,  Ann.  Ent.  Soc.  Am.,  v.  6,  p.  491. 

The  genus  Cornaphis  was  erected  by  Gillette  for  his  species  Corna- 
pJiis  populi.  In  his  description  it  is  stated  that  the  genus  is  closely 
related  to  Asiphum.  In  Cornaphis,  however,  there  are  large  corni- 
cles in  the  alate  form,  and  in  other  respects  it  seems  that  the  genus  is 
closely  related  to  Pachypappella.  In  that  genus,  however,  the 
media  is  once  forked,  whereas  in  Cornaphis  it  appears  to  be  simple,  at 
least  as  a  rule.  This  difference  has  led  the  writer  to  retain  a  genus 
with  lactea  as  type  rather  than  to  place  that  species  and  similar 
ones  in  Cornaphis. 

Characters. — Cornicles  present;  stem  mother  with  five-segmented  antenna?  and 
without  wax  plates.  Alate  form  with  six-segmented  antennae  armed  with  rather 
narrow  sensoria;  permanent  sensoria  ciliate.  Fore  wings  with  the  media  simple,  hind 
wings  with  both  media  and  cubitus  present;  wax  plates  present  in  the  apterous  form; 
sexes  small,  apterous  and  beakless;  the  oviparous  female  developing  only  one  egg. 

Forms  living  in  galls,  the  stem  mother  and  the  following  forms  living  in  the  same 
gall. 

Type  (monotypical),  Cornaphis  populi  Gill. 

Genus  DRYOPEIA  Kirkaldy. 
Plate  X,  U-Y. 

1857.  Endeis  Koch,  Die  Pflanzcnliiusc  Aphidon,  p.  312. 

1889.  Eudcis  Ashmead,  Ent.  Amcr.,  v.  5,  p.  189. 

1901.  Dryopeia  Kirkaldy,  The  Entomologist,  v.  37,  p.  279. 

1917.  Watabura  Matsumura,  Synopsis  of  the  Pemphigidae  of  Japan,  p.  89. 

In  1857  Koch  erected  his  genus  Endeis  with  two  species,  ~bdla 
Koch  and  rorea  Koch.  This  name  was  replaced  by  Dryopeia  in  1904 
by  Kirkaldy,  and  bella  has  been  definitely  placed  as  the  type. 


70  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

In  some  respects  the  genus  is  suggestive  of  Anoecia,  although  it 
seems  to  be  undoubtedly  a  Pemphiginid  and  will  no  doubt  be  so 
proven  by  the  sexual  forms. 

In  1917  Matsumura  erected  his  genus  Watabura  with  Watabura 
nisJiiyae  Mats,  as  type.  This  species  was  stated  to  resemble  a  Pem- 
phigus, excepting  that  the  antennal  segments  are  somewhat  different 
and  only  one  oblique  vein  is  in  the  hind  wing.  (Two  obliques  are 
shown  in  his  PI.  XII,  9).  The  antennae  are  armed  with  narrow  trans- 
verse sensoria  and  there  seems  little  doubt  that  this  genus  is  a  syno- 
nym of  Dryopeia.  It  is  noteworthy  that  the  life  history  of  the  type 
species  is  not  known,  but  it  is  thought  to  live  on  the  roots  of  trees. 
The  type  of  the  genus  Dryopeia  is  a  root  feeder. 

Characters. — Cornicles  present,  situated  on  broad  shallow  cones,  suggestive  of  those 
of  Anoecia.  Stem  mother  unknown,  apterous  forms  with  six-segmented  antennae. 
Alate  forms  with  six-segmented  antennae,  armed  with  narrow  transverse  sensoria. 
Fore  wings  with  media  simple,  hind  wings  with  one  oblique  vein.  Summer  forms 
subterranean,  living  on  the  roots  of  plants.  Spring  forms  and  sexes  unknown.  Apter- 
ous tarsi  one-segmented. 

Type  (fixed  by  Kirkaldy,  1906),  Endeis  bella  Koch. 

Genus  MORDWILKOJA  Del  Guercfo. 

Plate  XI,  A-G. 
1909.     Afordwilkoja  Del  Guercio,  Rivista  Patol.  Veget.,  v.  4,  p.  11. 

This  genus  was  erected  in  1909  for  the  peculiar  species  Byrsocrypta 
vagabunda  Walsh.  This  differs  in  the  antennae  quite  remarkably 
from  all  of  the  other  species  belonging  to  this  tribe.  The  difference 
is  in  the  long  unguis  of  the  sixth  segment.  However,  the  other  char- 
acters and  the  four-segmented  nature  of  the  antennae  of  the  stem 
mother  seem  to  place  it  with  little  doubt  in  the  Pemphigini. 

There  has  been  some  doubt  cast  by  Oestlund  on  the  determination 
of  Walsh's  species  and  this  has  led  Cockerell  to  propose  the  name 
oesflundi  for  the  species  now  known  so  well,  but,  as  Gillette  has 
pointed  out,  Walsh  evidently  accepted  the  insect  of  Riley  and  Monell 
as  the  same  species  as  his  vagabunda.  The  insects  Riley  had  were 
undoubtedly  the  species  we  know  and  the  writer  therefore  accepts 
vagabunda  and  the  generic  name  Mordwilkoja.  The  genus  was 
erected  with  the  name  vagabunda  used  as  type  and  not  oestlundi. 

Characters. — Cornicles  present  as  somewhat  elevated  rings.  Stem  mother  with  four- 
segmented  antennae,  the  unguis  of  segment  VI  slender  and  Aphis-like.  Permanent 
sensoria  ciliate.  Alate  form  with  five-segmented  antennae  which  are  armed  with 
narrow  transverse  sensoria.  Fore  wings  with  the  media  simple,  hind  wings  with  both 
media  and  cubitus  present. 

Forms  living  in  galls;  the  stem  mother  and  her  offspring  living  in  the  same  gall,  the 
alate  forms  leaving  the  galls  in  spring  or  early  summer.  Sexes  unknown,  but  no 
doubt  small,  apterous,  and  beakless. 

Type  (monotypical),  Byrsocrypta  vagabunda  Walsh. 


GENERIC   CLASSIFICATION  OF  APHIDIDAE.  71 

Genus  PACHYPAPPELLA,  n.  n. 

Plate  XI,  H-M. 
1909.     Pachypappa  Tullgren,  Arkiv  for  Zoologi,  Bd.  5,  no.  14,  p.  69. 

In  1854  the  genus  Pachypappa  was  erected  by  Koch,  with  marsu- 
pialis  and  vesicalis  in  the  genus.  A  study  of  marsupialis  shows  that 
this  species  is  in  reality  a  Pemphigus  as  it  shows  all  the  characters  of 
this  genus.  Tullgren,  1909,  noted  this  and  therefore  interpreted  the 
genus  differently.  Marsupialis  had,  however,  been  set  as  the 
type  of  the  genus.  Pachypappa  Koch,  therefore,  becomes  a  syno- 
nym of  Pemphigus,  and  Pachypappa  Tullgren  must  receive  a  new 
name  for  which  Pachypappella  is  here  given. 

Cfiaracters. — Stem  mother  without  cornicles  but  -with  wax  plates;  antennae  five- 
segmented.  Alate  form  with  cornicles;  antennae  six-segmented  and  with  transverse 
sensoria.  Fore  wings  with  media  once  branched,  hind  wings  with  both  media  and 
cubitua  present. 

Type  (present  designation),  Pachypappa  lactea  Tullgren. 

Genus  PEMPHIGUS  Harttg. 
Plate  XI,  N-U. 

1837.  Pemphigus  Hartig,  Jahresb.  u.  d.  Fortschr.  d.  Forstwiss.  und  forstl.  Naturk.,  v.  1,  p.  645. 

1839.  Byrsocrypta  Haliday,  Ann.  Nat.  Hist.,  v.  2,  p.  190. 

1840.  Brysocrypta  Westwood,  Int.  Mod.  Class.  Ins.,  Synopsis,  v.  2,  p.  118. 
1847.  Aphioides  Rondani,  Nuovi  Annali  Sci.  Nat.  Bologna  (2),  v.  8,  p.  439. 
1857.  A  myda  Koch,  Die  Pflanz.  Aphiden,  p.  301. 

1857.  Pachypappa  Koch,  Die  Pflanz.  Aphiden,  p.  263. 

1857.  Rhizomaria  Hartig,  Verhandl.  d.  Hils-Solling-Forstvereins,  Jahrg.  1856,  p.  52. 

1859.  Tychea  Koch,  Die  Pflanzenlatise  Aphiden,  p.  296. 

1885.  Kessleria  Lichtenstein,  Mon.  Puceron  du  PeupL,  p.  16. 

1904.  Hamadryaphis  Kirkaldy,  The  Entomologist,  v.  37,  p.  279. 

In  1837  Hartig  erected  his  genus  Pemphigus,  although  it  was  not 
until  1841  that  his  reference  to  the  genus  as  generally  cited  appeared. 
Passerini  in  1860  set  bursarius  as  type.  In  1839  Haliday  used  the 
generic  term  Byrsocrypta  but  mentioned  no  species.  In  1840  West- 
wood  referred  to  this  genus  as  Brysocrypta  and  gave  bursaria  L. 
as  type.  In  1859  Koch  erected  the  genus  Tychea  with  graminis 
Koch  as  type  (mono typical).  Schouteden  (1906)  has  described  the 
winged  form  of  Tychea  graminis  Koch  and  stated  that  it  is  a  typical 
Byrsocrypta.  The  writer  has  had  no  opportunity  to  study  specimens 
but  on  the  strength  of  this  statement  of  Schouteden  places  Tychea 
as  a  synonym  of  Pemphigus.  It  is  worthy  of  note,  however,  that 
Schouteden  did  not  mention  the  cornicles,  and  this  is  a  point  of 
considerable  difference  if  graminis  is  a  Pemphigus  or  if  it  belongs  to 
the  Fordini. 

In  1857  Hartig  described  the-  genus  Rhizomaria  with  piceae  Hartig 
as  type.  This  species,  however,  appears  to  be  a  typical  Pemphigus 
and  Rhizomaria  will  become  ~a  synonym. 

In  1857  Koch  erected  the  genus  Amycla  and  of  this  genus  fuscifrons 
Koch  has  boon  made  the  type.  The  writer  has  been  unable  to  obtain 
specimens  of  this  species  but  from  the  descriptions  it  seems  almost 


72  BULLETIN   826,   U.    S.   DEPARTMENT   OF   AGRICULTURE. 

certain  that  this  species  is  a  true  Pemphigus.  This  will  thus  make 
the  genus  Amycla  a  synonym. 

In  1847  Rondani  described  the  genus  Aphioides  of  which  bursaria 
Fab.  was  indicated  as  the  type  and  Aphioides,  therefore,  is  a  synonym. 

In  1854  Koch  erected  the  genus  Pachypappa  of  which  marsupialis 
Koch  has  been  made  the  type.  Marsupialis,  however,  is  a  typical 
Pemphigus.  Pachypappa  Koch,  therefore,  must  become  a  synonym. 
Tullgren  (1909)  used  Pachypappa  in  a  different  sense,  v  but  this  is 
discussed  under  the  genus  Pachypappella. 

In  1886  Lichtenstein  erected  the  genus  Kessleria  for  spiroihica 
and  this  name  was  replaced  by  Hamadryaphis  Kirk,  in  1904.  A 
study  of  this  species,  however,  shows  that  it  is  a  typical  Pemphigus. 
Therefore,  these  two  names  will  become  synonyms. 

Characters. — Cornicles  present;  wax  plates,  if  present,  weakly  developed;  stem- 
mother  with  four-segmented  antennae ;  alate  form  with  six-segmented  antennae  which 
are  armed  with  narrow,  oval  or  somewhat  irregular  sensoria.  Fore  wings  with  the 
media  simple;  hind  wings  with  both  media  and  cubitus  present  Sexes  small,  apter- 
ous, and  beakless.  Oviparous  female  developing  only  one  egg. 

Forms  living  in  galls,  the  stem-mother  and  her  offspring  in  the  same  gall,  the  alate 
forms  typically  leaving  the  galls  in  the  spring. 

Type  (fixed  by  Passerini,  1860),  Aphis  bursaria  L. 

Genus  PHLOEOMYZUS  Horvath. 
Plate  XI,  W-BB. 

1886.    Lowia  Lichtenstein,  Mon.  Puceron  Peupl.,  p.  37. 
1896.    Phloeomyzus  Ilorvath,  Wien.  Ent.  Zeit.,  v.  15,  p.  5. 

In  1886  Lichtenstein  erected  the  genus  Lowia  with  Schizoneura 
passerinii  Sig.  as  type  but  as  this  name  had  been  used  previously  it 
was  replaced  in  1896  by  Phloeomyzus  Horvath. 

It  is  with  some  hesitation  that  the  writer  places  this  genus  in  the 
Pemphigini,  In  some  respects  it  suggests  the  Melaphini,  while 
in  many  respects  it  strongly  suggests  the  Thelaxini  or  even  the 
Phyllaphidina.  Indeed,  to  the  Melaphini  it  shows  striking 
resemblances.  Without  a  study  of  the  sexual  forms  it  will  be  very 
difficult  to  place  the  genus  definitely.  All  that  can  be  done  at  the 
present  time  is  to  place  it  tentatively  with  the  forms  with  which  it 
appears  to  be  related,  and  if  further  study  shows  this  to  be  incorrect 
the  genus  can  be  placed  definitely  with  its  allies. 

Characters.— Cornicles  present,  very  slightly  elevated.  Apterous  form  with  six- 
segmented  antennse.  Alate  form  with  six-segmented  antennae  which  are  rather 
slender  and  without  secondary  sensoria.  Fore  wings  with  the  media  once  branched, 
hind  wings  with  both  media  and  cubitus  present.  Large  wax  plates  present  on  the 
abdomen.  Wings  held  flat  in  repose. 

Forms  living  free  upon  the  bark  of  trees  in  colonies. 

Type  (monotypical),  Schizoneura  passerinii  Sig. 

Study  based  on  specimens  received  from  Mordwilko  from  Warsaw, 
Poland,  and  notes  by  Pergande  on  type  specimens  loaned  by  Horvath. 


GENERIC   CLASSIFICATION   OF  APHIDIDAE.  73 

Genus  RHIZOCTONUS  Mokrzecky. 
1895.    Rhizoctonus  Mokrzecky,  Horae.  Soc.  Ent.  Ross.,  v.  30,  p.  438. 

The  genus  Rhizoctonus  was  erected  for  ampelinus  Mok.,  a  species 
occurring  on  the  vine.  Through  the  kindness  of  H.  F.  Wilson  the 
writer  has  been  able  to  examine  a  slide  containing  alate  forms. 
These,  however,  are  in  a  very  poor  condition  and  it  is  impossible  to 
determine  whether  or  not  cornicles  are  present.  The  antennae  too 
are  much  distorted.  This  slide  seems  to  indicate,  however,  that 
ampelinus  is  somewhat  related  to  passeriniiSig.,  an  hypothesis  which 
is  strengthened  by  the  fact  that  both  species  hold  the  wings  flat 
in  repose.  The  genus,  therefore,  is  placed  here  with  some  hesitation. 

Characters. — Antennae  of  six  segments,  without  secondary  sensoria  and  rather  thick. 
Fore  wings  with  the  media  simple.  Hind  wings  with  both  media  and  cubitus  present. 
Wings  held  flat  in  repose.  Cauda  and  anal  plate  rounded. 

Type  ("monotypical),  Rhizoctonus  ampelinus  Mok. 

Tribe  MELAPHINI. 

The  tribe  Melaphini  is  suggestive  both  of  the  Pemphigini  and  of 
the  Eriosomatini.  It  is,  however,  quite  distinct  from  both.  The 
habits  more  nearly  resemble  those  of  the  Pemphigini.  The  tribe 
is  placed  here  but  a  study  of  the  sexes  may  show  that  it  really  belongs 
to  the  Hormaphidinae.  The  typical  host  group  is  Rhus,  and  the 
galls  are  typically  fall  galls. 

Characters. — Gall-inhabiting  forms.  Cornicles  absent;  antennae  of  the  alate  form 
of  five  or  six  segments  armed  with  somewhat  oval  or  linear  transverse  sensoria.  Sexual 
forms  not  known. 

The  genera  may  be  separated  as  follows: 

KEY  TO  THE  GENERA  OF  THE  MELAPHINI. 

1 .  Both  media  and  cubitus  present  in  the  hind  wings 2. 

Only  the  media  present  in  the  hind  wings Aploneura. 

2.  Stigma  of  four  wings  pointed  on  distal  portion  and  extending  some  distance 

Melaphis. 

Stigma  stopping  abruptly  on  distal  extremity 3. 

?>.     Antennae  of  five  segments Nurudea. 

Antenme  of  six  segments Pemphigella. 

Genus  APLONEURA. 

Plato  XII,  A-E. 

1S03.    A  plonciiri  Paswrini,  Aphididac  Italicao,  p.  78. 

I»i9.      Tttrcncma  Dcrbes,  Ann.  des  Sc.  Nat.  Zool.  (.">),  v.  11,  p.  106. 

181S.    Baizongia  Rondani,  Nuovi  Annali  dcllo  Sdenze  Natural!,  v.  9,  p.  35. 

The  genus  is  distinguished  quite  easily  from  related  one's  by  the 
venation  of  the  hind  wings,  the  relation  of  the  cubitus  and  anal  of  the 
fore  wings,  and  by  the  structure  of  the  antennae  The  positive  determi- 
nation of  the  insect  of  Fabricius  may  cause  Aploneura  to  fall  for 
Rondani's  name. 


74  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

Characters. — Cornicles  absent.  Stem-mother  with  five-segmented  antennae;  alate 
form  with  the  media  simple,  thecubitusand  anal  joined  near  their  bases.  Hind  wings 
with  only  the  media  present,  antennae  of  six  segments,  armed  with  large  subcircular 
or  elongate  sensoria.  Forms  living  in  true  galls. 

Type  (monotypical),   Tetraneura  lentici  Pass. 

Genus  MEL  APHIS  Walsh. 
Plate  XII,  F-K. 

1866.     Melaphis  Walsh,  Proc.  Ent.  Soc.  Phlla.,  v.  6,  p.  281. 
1883.    Schlechtendalia  Liehtenstein,  Stett.  Ent.  Zeit.,  v.  44,  p.  240. 
1905.    Abamalekia  Del  Guercio,  Rodia,  v.  3,  p.  364. 

Walsh  erected  his  genus  for  rhois  Fitch,  a  species  forming  galls  on 
sumach  and  in  his  discussion  mentioned  the  Chinese  gall,  wonder- 
ing if  it  could  be  congeneric.  The  writer  has  recently  published  an 
account  of  the  Chinese  gall1  and  therein  placed  Schlechtendalia  Licht. 
as  a  synonym  of  Melaphis.  Del  Guercio's  genus  was  erected  with 
his  lazarewi  as  type  and  placed  as  a  thelaxine.  Although  the  writer 
has  never  obtained  specimens  of  this  species  he  is  of  the  opinion 
that  it  can  not  possibly  be  one  of  these  insects  and  that  it  evidently 
belongs  in  the  Melaphini  where  the  shape  of  the  stigma  would  imme- 
diately place  it  as  a  Melaphis. 

Characters. — Cornicles  absent.  Apterous  form  with  five-segmented  antennae. 
Alate  form  with  either  five  or  six  segmented  antennee  which  are  armed  with  some- 
what linear  or  oval  sensoria.  Fore  wings  with  the  media  simple,  although  sometimes 
slightly  forked,  distal  extremity  of  the  stigma  rather  long  drawn  out;  hind  wings  with 
both  media  and  cubitus  present;  abdomen  with  distinct  wax  plates.  Forms  living 
in  galls  from  which  the  alate  forms  escape  in  late  summer  or  fall. 

Type  (monotypical),  Byrsoerypta  rhois  Fitch. 

Genus  NURUDEA  Matsumura. 
Plate  XII,  L-Q. 

1917.    Wurudea  Matsumura,  Synopsis  of  the  Pemphigidae  of  Japan,  p.  65. 
1917.    Nurudeopsis  Matsumura,  Synopsis  of  the  Pemphigidae  of  Japan,  p.  67. 
1917.    Fushia  Matsumura,  Synopsis  of  the  Pemphigidae  of  Japan,  p.  70. 

Matsumura  erected  his  genus  Nurudea  for  his  Nurudea  Hbqfushi,  a 
species  somewhat  similar  to  a  species  of  Melaphis.  This  species  differs 
quite  markedly,  however,  in  the  form  of  the  stigma.  At  the  same 
time  he  erected  the  genus  Nurudeopsis  with  N.  sliiraii  as  type.  This 
species  differs  little  from  ibofusTii  excepting  in  the  proportions  of  the 
segments,  and  in  the  fact  that  the  cubitus  and  first  anal  are  some- 
what closer  togetner  at  the  base.  The  writer  is  of  the  opinion  that 
these  characters  are  not  sufficient  on  which  to  form  another  genus. 
In  the  same  work  also  he  erected  the  genus  Fushia  with  Fushia  rosea 
Mats,  as  type.  This  species  differs  somewhat  from  the  type  of  Nuru- 
dea but  the  writer  believes  that  there  are  not  differences  sufficient 
to  cause  this  to  be  considered  as  a  separate  genus.  The  antennae 

i  Baker,  A.  C.    On  the  Chinese  gall  (Aphididae— Horn.).    In  Ent.  News,  v.  28.  p.  385-393,  1917. 


GEISTERIC   CLASSIFICATION  OF  APHIDIDAE.  75 

are  more  slender  and  of  somewhat  different  proportions  and  the 
cubitus  and  anal  of  the  fore  wings  are  united  somewhat  at  base.  In 
other  genera,  however,  this  difference  between  species  is  to  be  found, 
and  it  seems  wisest  not  to  adopt  it  in  this  group  as  of  generic  im- 
portance. 

Characters. — Cornicles  absent.  Antennae  of  five  segments  armed  with  linear  or 
somewhat  oval  sensoiia.  Fore  wings  with  the  media  simple,  the  stigma  normal;  hind 
wings  with  both  media  and  cubitus  present.  Sexes  unknown.  Forms  making  galls 
upon  the  leaves  of  plants. 

Type  (fixed  by  Matsumura,  1917),  Nurudea  ibofushi  Mats. 

Genus  PEMPHIGELLA  Tullgren. 

1909.     Pemphigclla  Tullgren,  Arkiv  for  Zool.,  v.  5,  p.  171. 
191S.     Dasia  Van  der  Goot,  Mem.  Ind.  Mus.,  v.  6,  p.  152. 

Characters. — Cornicles  absent.  Antennae  of  six  segments  armed  with  oval  scnsoria. 
Hind  wings  with  both  media  and  cubitus  present.  Sexes  unknown.  Species  form- 
ing galls  on  plants.  Type  (monotypical),  Tetrancura  cornicularia  Pass. 

Tribe  PROCIPHILINI. 

The  tribe  Prociphilim  contains  forms  which  specialized  in  some  direc- 
tions more  than  did  the  Eriosomatini.  In  other  ways,  however,  they 
appear  to  be  more  primitive  than  certain  genera  of  that  tribe.  The 
cornicles  have  here  disappeared  altogether  and  largo  wax  areas  have 
replaced  them.  In  their  habits  of  gall  formation,  however,  the 
Eriosomatini  are  more  advanced  than  are  members  of  the  present 
tribe. 

Characters. — Forms  living  in  crumpled  or  twisted  leaves  or  in  a  somewhat  complete 
gall  caused  by  the  rolling  up  of  the  leaves  of  the  host.  Wax  plates  present;  cornicles 
absent;  antennae  of  stem  mother  of  five  segments;  those  of  the  alate  form  six  segmented 
and  armed  with  narrow,  transverse,  or  somewhat  broadly  oval  scnsoria. 

KEY  TO  THE  GENERA  OP  THE  PROCIPHILINI. 

] .     Media  once  forked,  stem  mother  usually  without  wax  plates Asiphum. 

Media  simple,  stem  mother  with  several  rows  of  wax  plates 2. 

L'.     Wax  plates  large,  those  on   the  thorax  well  developed,  stem  mother  and 

offspring  living  together 3. 

Wax  plates  not  well  developed,  stem  mother  usually  in  a  gall  by  herself .  .Thecabius. 
?,.     Hensoria    narrow,    linear,    ciliate Prociphilus. 

Hensoria  somewhat  oval,  nonciliate Neoprociphilus. 

Genus  ASIPHUM  Koch. 


Plate  XII,  K-X. 

1857.  Asiphum  Koch,  Die  Pflanzenlause  Aphidon,  p.  246. 
1859.  Type  fixation,  Gerstaecker,  Bcricht  for  1857,  p.  249. 
1903.  Type  fixation,  Kirkaldy,  Can.  Ent .,  v.  37,  p.  418. 


The  genus  Asiphum  was  erected  by  Koch  with  two  species,  populi 
Fab.  and  ligustrinellura  Koch.  lie  listed  De  Geer's  work  in  the 
literature  under  populi  Fab.  The  populi  of  Fabricius  proves  to  be 
the  tremulae  of  De  Gecr.  Only  the  species  ligustrinettwn  was  placed 


76  BULLETIN   826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

in  the  genus  by  Liechtenstein  (1885)  and  this  species  has  been  indicated 
as  type  of  the  genus  by  Kirkaldy  (1905).  Tremulae  De  Geer  is  a  well- 
known  species  but  the  writer  has  been  unable  to  obtain  ligustrineUum 
and,  in  fact,  has  been  unable  to  learn  anything  definite  in  regard  to 
the  species.  The  following  conception  of  the  genus,  therefore,  is 
based  upon  tremulae  De  Geer  in  view  of  the  fact  that  ligustrineUum 
appears  to  be  unknown,  and  since  Aphis  populi  Fab.  was  indicated 
by  Gerstaecker  in  1859. 

Characters. — Cornicles  absent;  wax  plates  present  in  the  alate  forms;  stem  mother 
with  five-segmented  antennae.  AJate  form  with  six-segmented  antennae  which  are 
armed  with  rather  narrow  transverse  sensoria.  Fore  wings  with  the  media  once  forked, 
hind  wings  with  both  media  and  cubitus  present.  Forms  living  in  the  somewhat 
crumpled  leaves  of  their  host. 

Type  (fixed  by  Gerstaecker,  1859),  Aphis  populi  Fab~ 

Genus  NEOPROCIPHILUS  Patch. 

Plate  Xin,  A-F. 
1912.     NeoprocipWus  Patch,  Bui.  Me.  Agr.  Expt.  Sta.,  no.  202,  p.  174. 

The  genus  Neoprociphilus  Patch  is  very  close  indeed  to  Prociphilusr 
the  characters  which  separate  it  being  the  somewhat  more  oval  or 
rounded  sensoria  and  the  fact  that  the  sensoria  are  not  ciliate.  How- 
ever, in  some  of  the  species  of  ProcipJiilus,  particularly  in  the  fall 
forms,  somewhat  oval  sensoria  are  met  with.  It  is  retained  doubt- 
fully. 

Characters. — Stem  mother  with  five-segmented  antennae.  Cornicles  absent,  large 
wax  plates  similar  to  those  of  Prociphilus  present.  Alate  form  with  six-segmented 
antennte  which  are  armed  with  oval  or  subcircular  nonciliate  sensoria.  Fore  wings 
with  the  media  simple,  hind  wings  with  both  media  and  cubitus  present.  Sexes  small, 
apterous,  and  beakless.  Oviparous  female  developing  only  one  egg. 

Forms  living  free  upon  their  host,  the  stem  mother  and  following  generations  in 
company. 

Type  (monotypical),  Pemphigus  uttenuatus  O.  S. 

Genus  PROCIPHILUS  Koch. 
Plate  XIII,  G-X. 

1857.  Prociplrilus  Koch,  Die  Pflanzcnlause,  p.  279. 

1857.  Stagonia  Koch,  Die  Pflanzenlause,  p.  284. 

1875.  Holzneria  Liechtenstein,  Bui.  Soc.  Ent.  Fr.  (5)  v.  5,  p.  LXXVI. 

1917.  Nishiyana  Hatsumura,  Synopsis  of  the  Pi-mphigidac  of  Japan,  p.  90. 

In  1857  Koch  erected  his  genus  Prociphilus  with  three  species: 
lumeliae  Schrank,  erraticus  Koch,  and  gnaphalii  Kalt.  Later  in 
the  work  (p.  284)  he  used  xylostei  De  Geer  as  the  type  of  the  genus 
Stagonia.  This  species,  xylostei,  is  in  all  respects  similar  to  bumeliae 
and  therefore  Stagonia  becomes  a  synonym  of  Prociphilus.  In  1875 
Lichtenstein  erected  the  genus  Holzneria  with  poschingeri  Holzner 
as  type.  Poschingeri  has  been  considered  by  many  authors  as  the 
alternate  form,  of  bumcliae.  In  such  case  Holzneria  must  necessarily 


GENERIC    CLASSIFICATION    OF   APHIDIDAE.  77 

be  a  synonym  of  Prociphilus.  Should  poschingeri,  however,  be 
proven  to  be  a  distinct  species  it  is  so  similar  in  all  regards  that 
Holzneria  must  remain  a  synonym. 

In  1917  Matsumura  erected  the  genus  Nishiyana  with  N.  aomori- 
ensis  Mats,  as  type,  placing  it  close  to  Prociphilus.  From  this  genus 
he  separated  it  because  of  the  absence  of  wax  plates  in  the  thorax 
and  the  somewhat  shorter  antennae.  It  must  be  borne  in  mind, 
however,  that  the  specimens  he  had  were  fall  migrants.  Fall  mi- 
grants of  several  species  of  Prociphilus  show  very  reduced  wax  plates, 
and  in  some  these  are  absent  altogether,  although  distinct  in  the 
spring  migrants.  It  is  believed  that  this  genus  is  in  reality  Proci- 
philus. 

Characters. — Cornicles  aosent,  wax  plates  present,  very  large  and  well  developed. 
Stem  mother  with  five-segmented  antennas.  Alate  form  with  six-segmented  antennae 
armed  with  narrow  transverse  sensoria;  secondary  sensoria  fringed.  Fore  wings  with 
the  media  simple,  hind  wings  with  both  media  and  cubitus  present.  Sexes  small, 
apterous,  and  beakless.  Oviparous  females  developing  only  one  egg.  Forms  living 
in  pseudogalls,  the  stem  mother  and  her  offspring  together. 

Type  (fixed  by  Gerstaecker,  1859),  Aphis  bumeliae  Schr. 

Genus  THECABIUS  Koch. 

Plato  XIII,  O-U. 

1857.     Thecabius  Koch,  Die  rflanzcnliiuse,  p.  294. 

1886.    Bucktonia.  Lichtenstein,  Mortogr.  d.  puccrons,  p.  16. 

The  genus  Thecabius  was  erected  by  Koch  in  1857  for  his  species 
populneus.  This  species  proves  to  be  a  synonym  of  Pemphigus 
affinis  Kalt.  In  1886 .  Lichtenstein  erected  the  genus  Bucktonia 
with  o$wis  Kalt.  as  type.  Bucktonia,  therefore,  becomes  a  synonym. 

Characters. — Cornicles  absent;  wax  plates  present  but  not  prominently  developed 
as  in  Prociphilus.  Stem  mother  with  five-segmented  antenna?  and  rather  narrow 
sensoria,  secondary  sensoria  not  fringed;  fore  wings  with  media  simple,  hind  wings 
with  both  media  and  cubitus  present.  Sexual  forms  small,  apterous,  and  beakless, 
oviparous  female  developing  only  one  egg. 

Forms  living  in  galls,  the  stem  mother  usually  living  in  a  gall  by  herself. 

Type  (mpnotypical),  Thecabius  populneus  'Koch  (=Pemphigus  affinis  Kalt.) 

This  genus  is  closely,  related  to  Prociphilus  and  it  is  with  some 
hesitation  that  the  writer  places  it  as  distinct.  Certain  species, 
such  as  patchii  Gillette,  which  are  undoubtedly  congeneric  with 
qffinis,  do  not  show  the  typical  life  habit  of  the  stem  mother  living 
in  a  gall  alone.  However,  the  character  of  the  sensoria  and  the 
undeveloped  nature  of  the  wax  glands  may  serve  to  distinguish  the 
genus. 

Tribe  FORDINI. 

Members  of  the  Fordini  are  specialized  subterranean  forms  mostly 
living  in  the  nests  of  ants.  The  aphids  excrete  honeydew,  in  return 
for  which  they  are  tended  carefully  by  these  insects.  The  apterous 


78  BULLETIN  826,   U.   S.   DEPARTMENT   OF   AGRICULTURE. 

forms  generally  are  of  a  yellowish  or  brownish-yellow  color,  some- 
times a  milk  white.  Wax-secreting  plates  are  present  but  they 
are  not  developed  to  the  same  extent  as  are  those  of  the  Prociphilini. 
The  cornicles  are  lost  entirely  and  the  region  where  these  usually 
occur  is  occupied  by  wax  plates.  In  some  species  the  wax  areas  are 
reduced.  Some  species  are  armed  with  fine  hairs,  whereas  others 
are  almost  entirely  smooth.  The  eyes  in  the  apterous  forms  are 
composed  of  three  facets.  The  alate  forms  have  rather  short,  thick 
antennae  with  somewhat  oval  sensoria.  Three  genera  compose  the 
tribe  and  these  may  be  separated  as  follows: 

KEY  TO  THE  GENERA  OF  THE  FORDINI. 

1.  Antennse  of  the  alate  form  composed  of  five  segments Forda. 

Antennae  of  the  alate  form  composed  of  six  segments 2. 

2.  Sensoria  of  the  antennae  of  the  alate  form  small  and  scattered  over  moat 

of  the  segment,  a  central  triangular  wax  plate  on  the  thorax;  apterous 

form  with  six-segmented  antennas .Paracletua. 

Sensoria  of  the  antennae  of  the  alate  form  larger  and  more  evenly  placed;  the 
sensoria  sometimes  extending  evenly  across  the  segment.  Apterous  form 
with  five-segmented  antennae,  sometimes  with  six  segments  present Geoica. 

Genus  FORDA  Heyden 
Plate  XIII,  V-AA. 

1837.  Forda  Heyden,  Mus.  Sinkbg.,  v.  2,  p.  291. 

1841.  Rhizoterus  Hartig.  Zeit.  Ent.,  v.  3,  p.  363. 

1849.  Smynthurodes  Westwood,  Gardener's  Chron.,  p.  420. 

1896.  Pentaphis  Horvath,  Wien.  Ent.  Zeit.,  v.  15,  p.  2. 

1909.  Pentaphis  Del  Guercio,  Rivist.  Patol.  Vegetale,  n.  s.,  v.  3,  p.  332. 

1914.  Rectiwsus  Theobald,  The  Entomologist,  v.  47,  p.  28. 

In  1837  Heyden  erected  his  genus  Forda,  the  type  of  which  is 
formicaria  Heyden.  In  1896  Horvath  erected  his  genus  Pentaphis 
with  marginata  Koch  as  a  type,  while  Del  Guercio  in  1909  used 
trivialis  Pass,  as  the  type  of  a  genus  of  the  same  name.  Specimens 
of  marginata  Koch  from  Horvath  prove  that  this  species  in  every 
respect  is  similar  to  the  type  of  the  genus.  Pentaphis,  therefore,  will 
become  a  synonym  of  Forda.  Likewise  specimens  of  trivialis  show 
that  this  species  belongs  in  the  same  genus.  In  1841  Hartig  erected 
the  genus  Rhizoterus,  the  type  of  which  is  vacca.  According  to 
Lichtenstein  this  species  is  a  synonym  of  formicaria  Heyden,  and 
Rhizoterus  also,  then,  becomes  a  synonym. 

In  1914  Theobald  erected  his  genus  Rectmasus  with  his  buxtoni 
as  type.  He  based  his  genus  on  the  proportions  of  the  antennal 
segments,  their  length,  and  the  length  of  the  beak.  The  writei  is 
opposed  to  basing  genera  on  the  proportions  of  the  antennal  segments, 
far  in  species  in  which  these  are  of  different  proportions  a  very  close 
relationship  is  evident.  This  is  also  true  of  the  beak.  Many  Ameri- 
can species  .taken  in  ants'  nests  and  as  yet  undescribed  have  beaks 
ranging  from  small  to  longer  than  the  body,  but  they  are  all  evidently 


GENERIC   CLASSIFICATION   OF  APHIDIDAE.  79 

closely  related.  The  other  characters  mentioned  by  Theobald  are 
seen  to  be  present  in  the  type  species.  The  tubercles  he  figures  and 
describas  are  the  same  and  the  spines  on  the  first  and  second  antennal 
segments  are  evidently  the  thickened,  pointed,  chitinized  articula- 
tions of  the  segments  common  in  insects  of  this  type.  We  believe,1 
therefore,  that  Rectinasus  should  be  carried  as  a  synonym  of  Forda. 
Westwood's  genus  was  erected  on  his  betae  which  appears  to  belong 
here  as  recently  indicated  by  the  writer. 

Characters. — Cornicles  wanting;  apterous  forms  with  five-segmented  antennae  and 
eyes  of  three  facets.  Alate  form  with  five-segmented  antennae  and  medium-sized 
oval,  or  more  or  less  irregularly  shaped  sensoria.  Fore  wings  with  media  simple; 
hind  wings  with  both  media  and  cubitus  present,  arising  slightly  apart.  Subterranean 
forms  living  usually  in  the  nests  of  ants  and  tended  by  them. 

Type  (monotypical),  Forda  formicaria  Heyden. 

G«nua  GEOICA  Hart. 
Plate  XIV,  A-K. 

1894.  Qeoica  Hart,  18th  Kept.  State  Ent.  111.,  p.  101. 

I860.  Tychea  Passerini,  Gli  Afldi,  p.  30. 

1906.  Tychcoides  Schouteden,  Mem.  Soc.  Ent.  Belg.,  v.  12,  p.  194. 

1906.  Kaltenbachiclla  Schouteden,  Mem.  Soc.  Ent.  Belg.,  v.  12,  p.  194. 

1909.  Trifidaphis  Del  Guercio,  Rivista  Patol.  Vegetale,  n.  s.,  v.  3,  p.  332. 

1912.  Tullgrenia  V.  d.  Goot,  Tijdschr.  voor  Ent.,  v.  15,  p.  96. 

1913.  Trinacriclla  Del  Guercio,  Redia,  v.  9,  p.  169. 

1916.    Serrataphis  V.  d.  Goot,  Zur  Kenntniss  der  Blattlause  Java's,  p.  263. 

In  1860  Passerini  used  the  generic  name  Tychea  of  Koch  and  placed 
as  the  typical  species  pJiaseoli  Pass.  In  1863  he  used  the  name 
again,  listing  several  species  but  not  the  species  included  by  Koch. 
Therefore  his  interpretation  of  the  genus  can  not  be  correct.  In  1894 
Hart  erected  his  genus  Geoica  with  squamosa  as  type.  In  1906 
Schouteden  noticed  Passerini 's  mistake  and  suggested  the  name 
Tycheoides  but  made  cragrostidis  Pass,  the  type.  In  the  same  year 
he  erected  Kaltenbachiella  with  mentliae  Schout.  as  type.  In  the 
year  1909  Del  Guercio  erected  Trifidaphis  with  radicicola  Essig  as 
type.  In  1912  Van  der  Goot  noted  Passerini's  mistake  and  proposed 
the  name  Tullgrenia  for  the  Tychea  of  Passerini.  In  1916  Van  der 
Goot  erected  the  genus  Serrataphis  with  lucifuga  Zehntner  as  type. 

In  studying  cotypes  and  other  specimens  of  squamosa  certain 
generic  characters  are  evident.  The  species  is  subterranean.  It  has 
five-segmented  antennoe  in  the  apterous  form  and  six-segmented  ones 
in  the  alate.  It  is  true,  however,  that  the  apterous  form  sometimes 
has  only  four  segments  in  the  antennas  and  the  alate  five. 
Indeed,  in  some  alate  forms  there  is  a  five-segmented  antenna  on  one 
side  and  a  six-segmented  one  on  the  other.  One  of  these  five- 
segmented  antennae  was  figured  by  Hart.  One  wing  vein  only  was 
figured  in  the  hind  wing  by  Hart,  but  a  very  close  examination  shows 
that  both  the  media  and  cubitus,  though  faint,  are  present.  These 
are  very  difficult  to  trace  in  balsam  mounts.  In  giving  his  name 


80  BULLETIN   826,   U.   S.   DEPARTMENT  OF  AGRICULTURE. 

Tycheoides  Schouteden  makes  plain  that  he  is  naming  the  Tychea 
of  Passerini  and  yet  he  sets  a  different  type.  He  further  states: 
<'Le  genre  Tychea  est  vraisemblablement  destine"  a  desparaitre,  ses 
especes  appartenant  en  re'alite'  a  Tetraneura  ou  Byrsocrypta."  He 
evidently  is  speaking  here  of  the  Tychea  of  Koch,  since  specimens  of 
Tychea,  Pass,  do  not  possess  the  cornicles  of  either  of  the  two  genera 
mentioned. 

In  describing  Kaltenbachiella  Schouteden  gives  as  a  character  the 
four-segmented  antennae,  etc.,  of  the  apterous  form.  The  alate 
form  he  did  not  know,  but  from  his  description  of  the  pupa  it  evi- 
dently would  possess  six-segmented  antennae.  There  seems  little 
doubt  that  this  is  another  such  case  as  squamosa  where  the  apterous 
form  has  often  four-segmented  antennas,  although  five  is  the  normal 
number,  the  alate  form  normally  having  six.  Hart's  description 
has  led  Schouteden  astray  and  he  places  Geoica  close  to  Forda, 
separating  it  therefrom  by  the  venation  of  the  hind  wing.  Consider- 
ing all  of  these  facts  there  seems  little  doubt  that  Kaltenbachiella 
should  be  placed  as  a  synonym  of  Geoica. 

Specimens  of  pTiaseoli  show  very  similar  characters  in  every 
respect.  It  is  true  that  the  antennas  are  somewhat  longer  and  the 
hairs  simple,  but  in  every  respect  of  importance  the  insects  agree. 
The  apterous  form  has  five-segmented  antennae  and  the  alate  form 
has  six-segmented  ones  with  the  sensoria  very  similar  in  nature. 
The  cauda  also  is  very  similar.  It  is  evident  then  that  Tychea 
Pass,  and  Tullgrenia  V.  d.  Goot  become  synonyms  of  Geoica  Hart. 
Specimens  of  lucifuga  Zehntner  show  a  remarkable  resemblance  to 
squamosa  with  the  exception,  of  course,  of  the  squamae.  The  apterous 
form  has  five-segmented  and  the  alate  form  six-segmented  antennae. 
In  general  form  and  structure  of  the  caudal  extremity  the  insects  are 
the  same  arid,  therefore,  Serrataphis  V.  d.  Goot  will  become  a  syno- 
nym of  Geoica.  There  remains,  then,  to  discuss  the  genus  Trifidaphis 
Del  Guercio.  The  co types  of  the  type. species  show  a  close  resem- 
blance to  the  general  type  of  squamosa.  The  apterous  forms  have 
five-segmented  antennae  and  in  the  alate  form,  as  in  squamosa,  some 
forms  have  five  segments  and  some  forms  have  six.  The  general 
resemblance  in  other  respects  seems  to  prove  that  Trifidaphis  is  a 
synonym  of  Geoica. 

It  should  be  pointed  out  that  the  sexes  described  by  Hart  are  in  all 
probability  not  sexual  forms,  but  immature  specimens. 

In  1913  Del  Guercio  erected  the  genus  Trinacriella  for  his  new 
species  magnified.  He  gave  a  brief  description  stating  that  the 
apterous  forms  had  five-segmented  antennae  and  the  alate  forms  six. 
No  specimens  of  this  species  are  available  to  the  writer,  but  there 
seems  little  doubt  that  Trinacriella  will  become  a  synonym  of  Geoica. 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  81 

Characters.— Cornicles  Aranting;  apterous  form  usually  with  five-segmented  antennae 
and  eyes  of  three  facets.  With  the  intermediate  forms  more  facets  may  occur.  Alate 
form  with  usually  six-segmented  antennae  and  rather  large  oval  sensoria  with  distinct 
rims.  Fore  wings  with  media  simple.  Hind  wings  with  both  media  and  cubitus 
present,  though  these  may  be  faint  and  almost  obscured  in  balsam.  Cauda  large 
and  somewhat  rectangular  or  rounded.  Subterranean  forms  living  on  the  roots  of 
plants.  Sexes  small,  apterous,  and  beakless. 

Type  (monotypical),  Geoica  squamosa  Hart. 

Genus  PARACLETUS  Heyden. 

Plate  XIV,  L-S. 
1S37.    Paradetus  Heyden,  Mus.  Sinkbg.,  v.  2,  p.  295. 

The  genus  Paracletus  is  closely  related  to  Forda  Heyden  from  which 
it  may  be  distinguished  by  the  number  of  antennal  segments  in  both 
alate  and  apterous  forms.  As  with  other  genera  of  this  tribe  the  eyes 
of  the  apterous  form  consist  of  three  facets.  Intermediate  forms 
often  occur,  however,  in  which  the  intermediate  nature  is  indicated 
only  by  the  eyes  which  have  the  beginnings  of  compound  eyes,  such 
as  are  found  in  the  alate  form.  There  never  are,  however,  complete 
compound  eyes.  The  genus  was  erected  by  Heyden  in  1837.  The 
species  live  in  close  association  with  ants. 

Characters.—  Cornicles  absent.  Apterous  form  with  six-segmented  antennae  and 
eyes  of  three  facets;  alate  form  with  six-segmented  antennae  which  possess  many 
rather  small,  oval  sensoria.  Fore  wings  with  media  simple,  hind  wings  with  both  media 
and  cubitus  present,  arising  some  distance  apart.  Thorax  with  a  central  wax  plate. 
Forms  living  in  the  nest  of  ants  and  cared  for  by  them.  Sexes  small,  apterous,  and 
beakless.  In  some  cases  only  one  claw  is  met  with  on  the  foot,  while  in  other  cases 
the  normal  number  of  two  is  present.  This  appears  to  be  no  definite  character,  as 
sometimes  a  claw  is  dropped  from  one  foot  and  sometimes  from  another. 

Type  (monotypical),  Paracletus  cimiciformis  Heyden. 

Subfamily  IV,  HORMAPHIDINAE. 

The  genera  placed  in  this  subfamily  have  usually  been  placed 
with  the  Eriosomatinae,  or  Pemphiginae,  as  it  has  been  sometimes 
called.  Mordwilko,  however,  placed  these  forms  as  his  third  tribe 
under  the  subfamily  Aphidinae  next  to  his  tribe  Callipterea. 
Something  can  be  said  in  favor  of  both  of  these  placings.  In 
the  first  instance,  the  species  in  general  form,  antennal  structure, 
and  habit  of  gall  formation  are  no  doubt  suggestive  of  the 
Eriosomatinae.  On  the  other  hand,  their  structure  in  regard  to 
cauda  and  anal  plate  is  very  like  the  Callipterina  and  the  sexual 
forms  appear  to  have  a  development  of  their  own,  although  they 
are  nearer  in  many  ways  to  the  Aphidinae  than  to  the  Eriosomatinae. 

It  is  the  author's  belief  that  these  forms  should  constitute  a  sepa- 
rate subfamily.  It  has  developed  the  habit  of  gall  formation 
and  the  sensory  characters  which  usually  accompany  it,  while  at 
the  same  time  it  has  retained  in  the  sexual  female  the  normal  develop- 

141613°— 20— Bull.  826 6 


82  BULLETIN   826,   U.    S.   DEPARTMENT   OF   AGRICULTURE. 

ment  of  the  ovaries  found  in  the  more  primitive  groups,  and  has 
retained  in  both  sexual  forms  the  beak  and  the  ability  to  feed. 
This  at  once  suggests  a  different  line  of  development  from  that 
taken  by  the  Eriosomatinae,  although  in  some  of  its  habits  the 
Hormaphidinae  agrees  with  that  subfamily.  In  other  lines,  however, 
marked  differences  are  met  with  here  and  one  of  the  most  striking 
of  these  is  development  of  aleyrodiform  generations,  which  remain 
stationary  upon  the  host.  Such  a  development  is  never  met  with  in 
the  Eriosomatinae,  although  the  sexual  forms  are  much  more 
specialized. 

Since  many  of  the  genera  of  the  Hormaphidinae  are  gall  formers, 
sensoria  very  similar  to  those  met  with  in  the  Eriosomatinae  are 
met  with  here  also.  Indeed  the  same  annular  sensoria  found  in 
the  Eriosomatini  are  even  more  pronounced  in  the  Hormaphidinae 
and  the  sensoria  on  the  wing  bases  are  prominent  and  often, 
numerous. 

The  cornicles  in  the  present  subfamily  are  sometimes  absent  or, 
as  is  usually  the  case,  reduced  to  mere  rings.  In  some  genera,  how- 
ever, they  may  be  elevated  slightly  on  broad  shallow  cones,  somewhat 
suggestive  of  those  of  Anoecia.  No  prominent  cornicles,  however', 
occur. 

In  the  wing  venation  there  is  often  a  considerable  reduction 
and  this  shows  also  the  specialized  nature  of  the  insects.  The 
venation  is  comparable  to  that  met  with  in  the  Eriosomatinae.  In 
the  fore  wings  the  media  is  either  simple  or  once  branched,  the  radial 
sector,  cubitus,  and  anal  are  present,  but  the  cubitus  and  anal  are 
often  fused  near  their  bases.  In  the  hind  wings  both  the  media  and 
cubitus  are  sometimes  present,  but  often  only  the  media  remains. 

Great  specialization  in  wax-producing  organs  occurs.  In  many 
of  the  forms  these  agree  with  the  ones  found  in  the  Pemphigini. 
In  certain  aleyrodiform  generations  and  in  some  sexual  forms 
agglomerate  glands  or  rather  groups  of  small  glands  are  seen.  These 
may  be  arranged  in  different  ways  and  often  are  placed  about  the 
margin  of  the  insect  so  that  it  possesses  a  distinct  lateral  fringe, 
;  very  like  that  of  an  aleyrodid.  In  fact  some  of  these  insects  on  this 
account  are  very  often  mistaken  for  aleyrodids. 

The  sexual  forms  are  often  quite  small  and  possess  large  wax- 
producing  areas.  Others  may  lack  these.  All,  however,  develop 
to  normal  adults. 

The  habit  of  gall  formation  is  very  marked  here.  Indeed,  some 
species  form,  galls  on  two  different  species  of  plants,  migrating 
L  between  the  two. 

Characters. — Aerial  forms  living  in  galls  or  sometimes  free   upon  the  host.     The 

!  mesothorax  in  many  forms  altered  so  that  its  divisions  are  more  or  less  unobservable, 

the  entire  mesothorax  often   showing  as  only  one  plate.     Scalelike  or  aleyrodiform 


GENERIC   CLASSIFICATION   OF  APHIDIDAE.  83 

generations  often  developed.  Cornicles  often  reduced  to  mere  ringlike  openings  or 
entirely  absent.  Sexual  forms  small  and  apterous  but  with  fully  developed  beaks. 
Oviparous  female  laying  several  eggs. 

KEY  TO   THE   TRIBES   OF   THE   HORMAPHIDINAE, 

1.  Aleyrodiform  generations  developed 2. 

Aleyrodiform  generations  not  developed OREGMINI. 

2.  Cornicles  absent;  insects  usually  gall  formers HORMAPHIDINI. 

Cornicles  usually  present;  insects  usually  not  gall  formers CERATAPHIDINI. 

Tribe  HORMAPHIDINI. 

Members  of  this  tribe  are  distinguished  easily  from  those  of  other 
tribes  in  that  the  cornicles  are  absent  and  aleyrodiform  generations 
are  developed.  These  remain  more  or  less  quiescent  upon  the 
foliage.  Some  different  forms  of  the  species  are  often  gall  pro- 
ducers. All  secrete  wax  from  special  pores.  Considerable  variation 
is  met  with  in  the  development  of  the  aleyrodiform  generations. 
Sensoria  of  the  alate  forms  are  usually  narrow  and  annular. 

Only  two  genera  are  so  far  recorded. 

KEY  TO  THE  GENERA  OF  THE  HORMAPHIDINI. 

Antennae  of  the  alate  forms  composed  of  three  segments,  hind  wings  with 
only  the  media  present  '. Hormaphis. 

Antennae  of  the  alate  forms  composed  of  five  segments,  hind  wings  with  both 
media  and  cubitus  present Hamamelistes. 

Genus  HAMAMELISTES  Shimer. 
Plate  XIV,  T-X. 

1867.    Hamamelistes  Shimer,  Trans.  Am.  Ent.  Soc.,  v.  1,  p.  284.  • 
1896.     Tetraphis  Horvath,  Wicn.  Ent.  Zeit.,  v.  15,  p.  6. 

Shimer  included  two  species  in  this  genus,  spinosus  Shimer  and 
cornu  Shimer.  The  latter  species,  as  suspected  by  him,  is  a  syno- 
nym of  hamamelidis  Fitch.  This  species  has  been  made  the  type 
of  Hormaphis. 

Characters. — Cornicles  absent.  Stem  mother  with  four-segmented  antennae.  Aley- 
rodiform generations  developed.  Alate  form  with  five-segmented  antennae  which 
are  armed  with  numerous  annular  sensoria.  Wings  held  flat  in  repose;  fore  wings 
with  the  media  simple;  hind  wings  with  both  media  and  cubitus  usually  present;  cauda 
knobbed,  anal  plate  bilobed;  wax -secreting  areas  abundantly  present  in  the 
apterous  forms.  Sexes  small  and  apterous  but  with  beaks  developed,  oviparous 
female  laying  several  eggs. 

Forms  living  in  galls  upon  the  leaves  or  scale-like  on  the  leaves  or  twigs. 

Type  (one  unquestioned  species),  Hamamelistes  spinosus  Shimer. 


84  BULLETIN   826,   U.    3.   DEPARTMENT   OF   AGRICULTURE. 

Genus  HORMAPHIS  Ostcn-Sacken. 

Plate  XIV,  Y-FF. 
1861.     Hormaphis  Osten-Sacken,  Stcttiner  Ent.  Zeit.,  p.  422. 

The  genus  Hormaphis  was  erected  by  Osten  Sacken  for  a  species 
he  described  as  hamamelidis.  This  species  it  later  proved  was  the 
same  described  by  Fitch  as  Byrsocrypta  hamamelidis.  The  genus 
later  was  made  to  include  spinosus  Shimer,  but  the  distinction 
between  this  genus  and  the  one  described  by  Shimer  has  been 
pointed  out  by  Pergande.1 

Characters. — Cornicles  absent;  aleyrodiform  generations  developed;  wax  glands 
very  numerous;  stem  mother  with  three-segmented  antennae;  alate  forms  with  three- 
segmented  antennae  which  are  armed  with  distinct  annular  sensoria.  Wings  held 
flat  in  repose;  fore  wings  with  the  media  simple;  hind  wings  with  the  cubitus 
absent.  Sexiial  forms  small  and  apterous,  possessing  beaks,  oviparous  female  laying 
several  eggs. 

Type  (monotypical),  Hormaphis  hamamelidis  O.  S.  (Byrsocrypta  hamamelidis  Fitch). 

Tribe  OREGMINI. 

Characters. — Forms  living  in  galls  or  otherwise  upon  the  leaves  of  plants,  possess- 
ing cornicles  and  wax  secreting  glands.  Antennae  of  the  winged  forms  usually  armed 
with  annular  sensoria;  cauda  rounded  or  somewhat  knobbed,  anal  plate  somewhat 
bilobed.  No  aleyrodiform  generations  developed.  The  sexual  forms  appear  to  be 
unknown. 

KEY  TO  THE  GENERA  OF  THE  OREGMINI. 

1.  Vertex  with  two  horn-like  projections 2. 

Vertex  without  such  horn-like  projections 3. 

2.  Antennae  five-segmented Oregma. 

Antennae  four-segmented Ceratoglyphina. 

3.  Antennae  of  apterous  form  with  five  segments 4. 

Antennae  of  apterous  form  with  four  segments Glyphinaphis. 

4.  Media  of  fore  wings  once  branched Astegopteryx. 

Media  of  fore  wings  simple Mansakia. 

Genus  ASTEGOPTERYX  Karsch. 
Plate  XV,  Q-X. 

1890.    Asteyoptcryx  Karsch,  Bcr.  deutsch  Botan.  Ges.,  v.  8,  p.  52. 

1906.     Nipponaphis  Pcrgande,  Ent.  News,  v.  17,  p.  205. 

1916.    ScldzoneurapJiis  Van  der  Goot,  Zur  Kenntniss  dcr  Rlattlause  Java's,  p.  245. 

The  genus  Astegopteryx  was  erected  with  styracophila  Karsch 
as  type.  Another  species,  nekoashi,  was  described  by  Sasaki  before 
the  International  Congress  at  Brussels  in  1911  and  a  third  species, 
styraci,  was  described  by  Matsumura  in  1917.  It  is  possible,  there- 
fore, to  gain  a  fair  conception  of  the  characters.  In  1906  Pergande 
erected  his  genus  Nipponaphis  with  distychii  Perg.  as  type.  This 
species  was  stated  to  be  from  Distychium  racemosum  in  Japan,  on 

1  Pergandc,  T.    The  life  history  of  two  species  of  plant  lice  inhabiting  both  the  witch-hazel  and  birch. 
U.  S.  Dept.  Agr.  Bur.  Ent.  Tech.  Ser.  no.  9.    1901. 


GENERIC    CLASSIFICATION   OF"  APHIDIDAE.  85 

which  it  forms  galls.  In  1916  Van  der  Goot  erected  nis  Schizoneura- 
phis  with  gaTlorum  V.  d.  Goot  as  type.  This  species  was  said  to 
form  galls  on  Distylium  steUare. 

The  genus  Astegopteryx  can  be  separated  as  far  as  the  recognized 
forms  ure  concerned  by  the  proportion  of  the  antennal  segments 
and  some  variation  in  the  shape  of  the  stigma.  These  differences 
are  not,  however,  of  large  importance  and  Nipponaphis  should  be  a 
synonym  of  Astegopteryx.  In  the  same  way,  the  type  of  Van  der 
Goot's  genus  is  not  sufficiently  different  to  warrant  the  erection  of 
a  new  genus  and  Schizoneuraphis  also  should  be  considered  a  synonym. 

Characters. — Cornicles  broad  rings;  apterous  form  with  five-segmented  antennae; 
alate  form  with  five-segmented  antennae  which  are  armed  with  annular  sensoria. 
fore  wings  with  the  media  once  branched;  hind  wings  with  both  media  and  cubitua 
present.  Stigmal  vein  arising  rather  far  back  on  the  stigma.  Cauda  rounded,  anal 
plate  somewhat  bilobed;  forms  living  in  galls. 

Sexual  forms  unknown. 

Type  (mono typical),  Astegopteryx  styracophila  Karsch. 

Genus  CERATOGLYPHINA  Van  der  Geot. 

Tlate  XV,  M-P. 
1916.     Ceratoglyphina  Van  der  Goot,  Zur  Kenntniss  der  Blattlause  Java's,  p.  237. 

Characters. — Cornicles  present  as  mere  pores.  Vertex  with  two  hornlike  projec- 
tions. Antennae  of  four  segments;  cauda  and  anal  plate  both  rounded.  Winged 
orms  unknown . 

Type  (fixed  by  V.  d.  Goot,  1916),  Ceratoglyphina  bambusaeV.  d.  Goot 

Genus  GLYPHINAPHIS  Van  der  Goot. 

Plate  XV,  H-K. 

• 

1916.    Glyphinaphis  Van  der  Goot,  Zur  Kenntniss  der  Blattlause  Java's,  p.  232. 

Characters. — Cornicles  present  as  mere  pores;  antennae  of  four  segments  which  are 
armed  with  linear  sensoria.  Fore  wings  with  the  media  once  branched;  hind  wings 
with  both  media  and  cubitus  present;  cauda  knobbed;  anal  plate  rounded;  body 
covered  with  stout  hairs. 

Type  (fixed  by  V.  d.  Goot,  1916),  Glyphinaphis  bambusae  V.  d.  Goot. 

Genus  MANSAKIA  Matsumura. 

Plate  XV,  L. 
1017.     ^fantiakia  Mutsumura,  Synopsis  of  the  Pemphigidae  of  Japan,  p.  59. 

The  author  of  the  genus  Mansakia  stated  that  it  is  closely  allied 
to  Hormaphis,  but  it  would  appear  to  the  writer  to  be  related  to  the 
genera  in  the  Oregmini  as  understood  in  the  present  classification. 
The  presence  of  the  cornicles  would  indicate  that  the  genus  is  not 
related  as  closely  to  Hormaphis  as  to  Astegopteryx,  but  its  host  and 
the  nature  of  the  gall  would  place  it  close  to  Hamamelistes.  Since 
all  of  the  forms  are  not  known  it  is  impossible  to  state  positively 
its  position. 


86  BULLETIN    826,    U.    S.    DEPARTMENT   OF   AGRICULTURE. 

Characters. — Cornicles  present  as  mere  rings.  Antennae  of  five  segments  armed 
with  annular  sensoria.  Fore  wings  with  the  media  simple,  hind  wings  with  two  oblique 
veins.  Cauda  rounded,  anal  plate  somewhat  bilobed.  Sexes  not  known. 

Forms  causing  galls  upon  plants,  the  galls  usually  armed  with  numerous  long  pro- 
jections. 

Type  (fixed  by  Matsumura,  1917),  Mansalda  miyabei  Mats. 

Genus  OREGMA  Buckton. 
Plate  XV,  A-G. 

1893.    Oregma  Buckton,  Ind.  Mas.  Notes,  v.  3,  p.  87. 

1897.     Ceratovacuna  Zehntner,  Mededl.  Proefs.  Java,  n.  s.,  no.  37,  p.  29. 

In  1893  Buckton  established  his  genus  Oregma  with  'bambusae 
Buckton  as  type,  while  in  1897  Zehntner  established  his  Ceratovacuna 
with  lanigera  Zehntner  as  type.  Specimens  of  both  of  these  species 
sent  by  Zehntner  and  specimens  of  bambusae  from  Green  taken  in 
Ceylon  show  that  these  genera  must  be  considered  the  same. 

Characters. — Both  alate  and  apterous  forms  with  two  hornlike  projections  on  the 
vertex,  wax  gland  areas  present;  antennae  five-segmented,  those  of  the  alate  form 
with  narrow  annular  sensoria;  cauda  rounded  or  somewhat  knobbed;  fore'  wings  with 
media  twice  forked,  hind  wings  with  both  media  and  cubitus  present;  cornicles  broad, 
slightly  elevated  rings. 

Type  (monotypical),  Oregma  bambusae  Buckt. 

Tribe  CERATAPHIDINI. 

This  tribe  is  closely  related  to  the  Hormaphidini  but  differs  in  that 
very  distinct  cornicles  are  here  present.  The  apterous  forms  are 
scalelike  and  quiescent  and  feed  upon  the  surfaces  of  the  leaves. 
The  alate  forms  possess  annular  sensoria.  Wax  secretion  is  abundant. 

The  genera  may  be  separated  as  follows: 

KEY  TO  THE  GENEEA  OF  THE  CEKATAPHJDINI. 

Antennae  of  the  aleyrodiform  generations  of  five  segments Aleurodaphis. 

Antennae  of  the  aleyrodiform  generations  of  four  segments Cerataphis. 

Antennae  of  aleyrodiform  generations  of  three  segments Thoracaphis. 

Genus  ALETTRODAPHIS  Van  der  Goot. 

Plate  XVI,  A-E. 
1916.    Aleurodaphis  Van  der  Goot,  Zur  Kenntniss  der  Blattlause  Java's,  p.  239. 

This  genus  was  erected  for  one  species  occurring  in  Java. 

Characters. — Form  flat,  aleyrodiform,  three  distinct  -divisions  evident;  cornicles 
present  as  mere  rings.  Margin  with  wax  secreting  glands;  dorsum  also  with  many 
small  glands.  Cauda  rather  elongate  and  knobbed,  anal  plate  bilobed.  Antennae 
five-segmented;  eyes  of  the  apterous  forms  with  three  facets. 

Type  (fixed  by  Van  der  Goot,  1916),  Aleurodaphis  blumeae  V.  d.  Goot. 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  87 

Genus  CERATAPHIS  Lichtenstein. 
Plate  XVI,  F-M. 

1862.    Boisduvalia  Signoret,  Ann.  Ent.  Soc.  France  (4),  v.  8,  p.  400. 
1882.     CeratapJiis  Lichtenstein,  Bui.  Ent.  Soc.  France  (6),  v.  2,  p.  xvi. 

Signoret  erected  his  genus  Boisduvalia  in  connection  with  his 
aleyrodid  monograph.  He  placed  Coccus  lataniae  Bois.  as  type. 
Later,  however,  he  considered  this  a  coccid  genus.  The  name  was 
used  in  the  Diptera  in  1830,  and  is  therefore  not  available.  The 
name  Cerataphis  used  by  Lichtenstein  in  1882  appears  as  the  next 
name  applied  to  the  genus. 

Characters. — Cornicles  present  as  mere  rings.  Apterous  form  with  four-segmented 
antennae  and  aleyrodiform,  with  two  divisions  to  the  body;  wax  glands  prominent; 
vertex  with  two  hornlike  projections;  alate  form  with  five-segmented  antennae,  the 
segments  armed  with  narrow  annular  sensoria.  Fore  wings  with  the  media  once 
branched,  hind  wings  with  both  media  and  cubitus  present.  Cauda  knobbed;  anal 
plate  bilobed. 

Type  (monotypical),  Coccus  lataniae  Bois. 

Genus  THORACAPHIS  Van  der  Goot 

Plate  XVI,  N-W. 
1916.     Tharacaphis  Van  der  Goot,  Zur  Kenntniss  der  Blattlause  Java's,  p.  242. 

Only  the  apterous  form  of  one  species  of  this  genus  has  been 
described.  Other  species,  however,  are  available  for  study,  through 
the  generosity  of  Professor  Van  der  Goot. 

Characters. — Cornicles  present  and  quite  distinct,  occasionally  absent,  however, 
in  the  apterous  form.  Apterous  form  with  three-segmented  antennae,  flat  and  with  a 
posterior  lobe.  Alate  form  with  five-segmented  antennae  armed  with  annular  sen- 
soria. Fore  wings  with  the  media  once  branched;  hind  wings  with  both  media  and 
cubitus  present.  Cauda  somewhat  knobbed,  anal  plate  bilobed. 

Type  (monotypical),  Thoracaphis  arboris  V.  d.  Goot. 


88  BULLETIN   826,   U.    S.    DEPARTMENT  OF   AGRICULTURE. 

GENERA  NOT  PLACED. 

A  number  of  genera  have  been  described  which  the  writer  has 
been  unable  to  place.  These  genera  are  discussed  in  the  following 
notes. 

Genus  RHIZOBIUS  Burmeister. 

1S35.  Rhizobius  Burmeister,  Handbuch  der  Entomologic,  p.  78. 

1819.  Rhizophlhiridum  Van  der  Hoeven,  Handb.  Dierkunde  v.  1,  p.  508. 

1830.  Rhyzoicus  Passerini,  Gli  Afldi,  p.  30. 

1863.  Rizolrius  Passerini,  Aphididae  Italicae,  p.  79. 

1919.  Rhizoicus  Del  Guercio,  Redia,  v.  12,  p.  251. 

The  genus  Rhizobius  has  generally  been  considered  as  a  good 
aphid  genus  and  writers  have  referred  to  species  in  this  genus  as 
having  but  one  claw  to  the  tarsus.  However,  as  indicated  under 
Paracletus  the  writer  believes  this  is  a  variable  character  and  we 
have  no  definite  knowledge  in  regard  to  piloseUae  Burm.  Buckton's 
species  of  course  was  not  in  the  original  genus  and  therefore  can  not 
be  used  as  type  nor  was  it  in  Passerini's  conception  of  Rhyzoicus. 
After  placing  Rhyzoicus  Pass,  with  jujubae  Buckton  as  type,  Del 
Guercio  erects  the  genus  Neorhizobius,  distinguished  by  having  two 
claws,  and  in  which  he  places  graminis  Thos.,  poae  Del  Guercio, 
stramineus  Del  Guercio,  and  ulmipTiilus  Del  Guercio. 

In  1860  Passerini  set  soncTii  Pass,  as  the  type  of  Rhizobius  Burm., 
and  in  a  footnote  suggested  the  name  Rhyzoicus  as  a  new  name  for 
Rhizobius,  since  this  name  had  previously  been  used  in  the  Coleoptera. 
Such  procedure,  however,  is  not  allowable  since  sonchi  Pass,  was  not 
in  the  original  genus.  Of  the  two  species  in  the  original  genus 
pilosettae  Burm.  has  been  accepted  as  type. 

Del  Guercio  in  1917  used  the  generic  name  Rhizoicus  Pass., 
spelling  it  with  an  "i"  instead  of  a  "y",  and  jujubae  Buckton  as 
the  type. 

In  the  writer's  opinion  the  genus  Rhizobius  must  remain  unknown 
until  the  type  species  pilosellae  becomes  known  and  carefully  studied. 

The  name  Rhizophthiridum  was  given  to  this  genus  to  replace 
Rhizobius  Burm. 

Genus  NEORHIZOBIUS  Del  Guercio. 
1917.     Neorhizobius  Del  Guercio,  Redia,  v.  12,  p.  251. 

As  indicated  under  the  discussion  of  Rhizobius,  four  species  are 
placed  in  this  genus  by  Del  Guercio.  Three  species  are  described 
as  new  and  only  in  the  apterous  forms. 

Two  of  these  forms  have  five-segmented  antennae  and  one  of  them 
four-segmented  ones,  and  when  the  alate  forms  are  found  they  will 
in  all  probability  be  shown  to  represent  species  either  of  Forda  or 
Geoica.  The  basing  of  genera  upon  the  relative  lengths  of  the 
antennal  segments  would  create  a  very  large  number  of  genera  and 
separate  related  forms.  The  genus  Neorhizobius,  therefore,  the 


GENERIC    CLASSIFICATION   OF   APHIDIDAE.  89 

writer  considers  composed  of  apterous  forms,  which  really  belong  in 
older  and  well-recognized  genera,  but  which  can  not  be  definitely 
placed  until  the  alate  forms  have  been  secured  and  studied. 

Genus  SCHOUTEDENIA  Riibsaamen. 
1905.    Schautedcnia  Riibsaamen,  Marcellia,  v.  4,  p.  19. 

This  genus,  which  was  described  for  ralumensis  Rub.,  is  here  listed 
as  unknown.  The  gall  formed  by  the  species  is  described  but  the 
writer,  never  having  been  able  to  obtain  either  gall  or  insect,  would 
be  able  only  to  guess  at  its  position  from  the  description. 

Genus  CLAVIGERUS  Szfepligeti. 
Clavigerus  Szdpligeti,  Kovar&szati  Lapok,  v.  1,  p.  4. 

This  genus  was  described  in  the  only  volume  issued  of  the  Journal 
cited.  The  writer  has  been  unable  to  secure  a  copy  or  any  details 
of  the  description  given. 

ADDENDA. 

The  following  generic  names  have  been  employed  by  Mordwilko 
(Fauna  Russ.  vol.  1,  Aphidodea)  without,  apparently,  the  mention 
of  any  species  in  connection  therewith.  They  have  not  been  con- 
sidered in  the  foregoing  paper:  Anameson,  Aorison,  Chaetosiphon, 
Corylobium,  Elatobium,  Euaulax,  Eurythaphis,  Halmodaphis, 
Impatientinum,  Jaxart  aphis,  Nasonovia,  Orobion,  Paczoskia, 
Sitobium,  Staticobium,  Tlja,  Turanaphis,  Uroleucon  (subgenus), 
Uromelan.  The  subgenera  Dactynotus  Raf.,  Cladoxus  Raf.,  and 
Adactynus  Raf.,  have  not  been  considered. 


INDEX  TO  GENERA. 


Abamalekia  Del  Guercio 

Abura  Matsumura 

Acanthaphis  Matsumura... 
Acanthocallis  Matsumura. . . 

Acaudus  Van  der  Goot 

Acyrthosiphon  Mordwilko  . 

Akkaia  Takahashi 

Aleurodaphis  Van  der  Goot. 

Amphorophora  Buckton 

Amycla  Koch 

Anoecia  Koch 

Anomalaphis  Baker 

Anuraphis  Del  Guercio 


Page. 

74 

43 

54 

29 

41 

56 

53 

86 

54 

71 

13 

52 

42 

Aphioides  Passerini 36 

Aphis  Linnaeus 43 

Apholdes  llondani 71 

Aploneura  Passerini 73 

Arakawana  Matsumura 34 

Arctaphis  AValker ."  33 

Arimakla  Matsumura 43 

Arlstaphis  Kirkaldy 36 

Asiphum  Koch 75 

Asiphonaphis  Wilson  &  Davis 44 

Aspidaphis  Gillette 44 

Astegopteryx  Karsch 84 

Atarsos  G  illette 44 

Atheroides  Ilaliday 33 

Aulacorthum  Mordwilko 57 

Baizongia  Rondanl 73 

Boisduvalla  Signoret 87 

Brachycaudus  Van  der  Goot 42 

Brachycolus  Buckton 45 

Brachysiphum  Van  der  Goot 45 

Brachyunguis  Das 49 

Bradyaphis  Mordwilko 32 

Brevicoryne  Van  der  Goot 45 

Brysocrypta  Westwood 71 

Bucktonia  Lichtenstein 77 

Byrsocrypta  Haliday 71 

Byrsocrypta  Tullgren 67 

Calaphis  Walsh 26 

Callaphis  Walker 27 

Callipterinella  Van  der  Goot 26 

Callipteroidcs  Mordwilko 28 

Callipteroides  Van  der  Goot 28 

Callipterus  Koch 27 

Callipterus  Van  der  G  oot 29 

Capitophorus  Van  der  Goot 55 

Carolinaia  Wilson 45 

Cavariella  Del  Guercio 46 

Cerataphis  I/ichtenstein 87 

Ceratoglyphina  Van  der  Goot 85 

Ceratovacuna  Zehntner 86 

Cerosipha  Del  Guercio 4ii 

Cervaphis  Van  der  Goot 53 

Ceylonia  Buekton 51 

Chaitophorinella  Van  dt> r  Goot 34 


Page. 

Chaitophorus  Koch 33 

Chelymorpha  Clark 34 

Chromaphis  Walker 27 

Cinaria  Curtis 15 

Cladobius  Koch 36 

Clavigerus  Sze'pligeti 89 

Colopha  Monell 60 

Coloradoa  Wilson 49 

Cornaphis  Gillette 69 

Corynosiphon  Mordwilko 46 

Cryptosiphum  Buckton 46 

Dasia  Van  der  Goot 75 

Davisla  Del  Guercio 17 

Dentatus  Van  der  Goot 42 

Dielcysmura  Mordwilko 56 

Dilachnus  Baker 16 

Drepanaphis  Del  Guercio 31 

Drepaniella  Del  Guercio 57 

Drepanosiphum  Koch 32 

Dryaphis  Del  Guercio 18 

Dryobius  Koch 18 

Dryopeia  Kirkaldy 09 

Eichochaitophorus  Essig 33 

Endeis  Koch C9 

Eriosoma  Leach 6G 

Essigella  Del  Guercio 14 

Eucallipterus  Schouteden 28 

Euceraphis  Walker 28 

Eudeis  Ashmead 69 

Eulachnus  Del  G  uercio 15 

Eunectarosiphon  Del  Guercio 54 

Eutrichoslphum  Essig  &  Kuwana 38 

Forda  Heyden 78 

Francoa  Del  Guercio 55 

Fullawaya  Essig 37 

Fullawayella  Del  Guercio 59 

Fushia  Matsumura 74 

Geoica  Hart 79 

Georgia  Wilson 6" 

Glyphina  Koch 21 

Glyphinaphis  Van  der  Goot 85 

Gobaishia  Matsumura 67 

Greenidea  Schouteden 37 

Greenidea  Wilson 38 

Greenideoida  Van  der  Goot 38 

Hamadryaphis  Kirkaldy 71 

Hamamelistes  Shinier 83 

Hannabura  Matsumura 41 

Hayhurstia  Del  Guercio 47 

Heteroneura  Davis 47 

Holzneria  Lichtenstein 76 

Hormaphis  Osten  Sacken 84 

Hyadaphis  Kirkaldy 47 

Hyalopteroides  Theobald 56 

Hyalopterus  Koch 47 

Hysteroueura  Davis 47 

91 


92 


BULLETIN   826,    U.    S.    DEPARTMENT   OF   AGRICULTURE. 


Page. 

Idiopterus  Davis 60 

lllinoia  Wilson 56 

Kallistaphis  Kirkaldy 28 

Kaltenbachiella  Schouteden 79 

Kessleria  Lichtenstein 71 

Lachniclla  Del  Guercio 15 

Lachnus  Burmeister 15 

Liosomaphis  Walker 48 

Longicaudus  Van  dor  Goot 49 

Longistigma  Wilson 17 

Longiunguis  Van  der  Goot 43 

Lowia  Lichtenstein 72 

Loxerates  Rafmesquc 43 

Macchiatiella  Del  Guercio 41 

Macrosiphon  Del  Guercio 57 

Macrosiphoniella  Del  Guercio 50 

Macrosiphum  Del  Guercio 54 

Macrosiphura  Oestlund 54 

Macrosiphum  Fasserini 57 

Macrosiphum  Van  der  Goot 56 

Mansakia  Matsumura 85 

Mastopoda  Oestlund 48 

Megoura  Buckton 57 

Melanaphis  Van  der  Goot 43 

Melaphis  Walsh 74 

Melanoxantherium  Schouteden 36 

Melanoxanthus  Buckton 36 

Metaphis  Matsumura 46 

Metopeurum  Mordwilko 56 

Micrella  Essig -  -  -  -  33 

Micromyzus  Van  der  Goot 59 

Microparsus  Patch 00 

Microsiphon  Del  Guercio 43 

Microsiphum  Cholodkovsky 49 

Mimaphidus  Rondani 06 

Mindarus  Koch 62 

Monaphis  Walker 32 

Monellia  Oestlund 29 

Monvilkoja  Del  Guercio 70 

Myzaphis  Van  der  Goot 43 

Myzocallis  Passerini 29 

Myzodes  Mordwilko 57 

Myzoides  Van  der  Goot 57 

Myzopsis  Matsumura 57 

Myzoxylus  Blot 66 

Myzus  Passerini 57 

Nectarophora  Oestlund 57 

Nectarosiphon  Schouteden 54 

Neocallipterus  Van  dor  Goot 28 

Neomyzus  Van  der  Goot 57 

Neophyllaphis  Takahashi 24 

Neoprociphilus  Patch 76 

Neorhizobius  Del  Guercio 88 

Neosymydobius  Baker 32 

Neothomasia  Baker 35 

Neotoxoptera  Theobald 60 

Neotrama  Baker 20 

Nippocallis  Matsumura 27 

Nippolachnus  Matsumura 14 

Nipponaphis  Pergande 84 

Nipposiphum  Matsumura 46 

Nishiyana  Matsumura 76 

Nurudea  Matsumura 74 

Nurudeopsis  Matsumura 74 

Oedisiphum  Van  der  Goot 45 

Oregma  Buckton 


Page. 

Ovatus  Van  der  Goot 57 

Pachypappa  Koch 71 

Pachypappa  Tullgren 71 

Pachypappella  Baker 71 

Panaphis  Kirkaldy 27 

Paracletus  Heyden 81 

Patchia  Baker 34 

Pcmphigella  Tullgren 75 

Pemphigus  Hartig 71 

Pentalonia  Coquerel 61 

Pentaphis  De!  Guercio 78 

Pentaphis  Horvath 78 

Pcrgandeidia  Schouteden. 49 

Periphyllus  Van  der  Hoeven 34 

Phillophorus  Thornton 34 

Phloeomyzus  Horvath 72 

Phorodon  Passerini 58 

Phyllaphis  Koch 24 

Phymatosiphum  Davis 31 

Prociphilus  Koch 76 

Protolachnus  Theobald 15 

Protrama  Baker 19 

Pterocallis  Passerini 29 

Pterochlorides  Archangelsky 18 

Pterochlorus  Rondani 18 

Pterocpmma  Buckton 36 

Ptychodes  Buckton 27 

Rectinasus  Theobald = 78 

Rhizoberlesia  Del  Guercio 49 

Rhizobius  Burmeister 88 

Rhizoctonus  Mokrzecky 73 

Rhizoicus  Del  Guercio 88 

Rhizoicus  Passerini 88 

Rhizomaria  Hartig 71 

Rhizophthiridum  Van  der  Hooven 88 

Rhizoterus  Hartig 78 

Rhopalosiphum  Koch 49 

Rhopalosiphum  Passerini 57 

Rhopalosiphum  Van  der  Goot 54 

Rhopalosiphon  Scudder 49 

Rhopalosiphoninus  Baker 58 

Rhynchocles  Altum 18 

Rizobius  Passerini 88 

Saltusaphis  Theobald 31 

Sanbornia  Baker 50 

Sappaphis  Matsumura 42 

Schizodryobius  Van  der  Goot IS 

Schizolachnus  Mordwilko 16 

Schizoneura  Hartig 66 

Schizoneuraphis  Van  der  Goot 84 

Schlechtendalia  Lichtenstein 74 

Schoutedenia  Riibsaamen 89 

Semiaphis  Van  der  Goot 

Serrataphis  Van  der  Goot 79 

Setaphis  Van  der  Goot 39 

Shivaphis  Das 

Sipha  Passerini 

Siphocoryne  Passerini 47,49 

Siphonaphis  Van  der  Goot 49 

Siphoualrophia  Swain 

Siphonocallis  Del  Guercio 26 

Siphonophora  Koch 

Smynthurodes  Westv/oo  1 

Stagonia  Koch ' a 

Stenaphis  Del  Guercio 43 

Stephensonia  Das 49 


GENERIC    CLASSIFICATION    OF   APHIDIDAE. 


93 


Stomaphis  Walker  

Page. 
18 

Trifidaphis  Del  Guercio  

Page. 
79 

Subcallipterus  Mordwilko 

29 

Trinacriella  Del  Guercio 

79 

Symydobius  Mordwilko  

30 

Tuberculatus  Mordwilko  

29 

Takecallis  Matsumura  

29 

Tuberculoides  Van  der  Goot  

29 

Tamalia  Baker  

24 

Tuberodryobius  Das  

18 

Tetraneura  Hartig 

es 

Tuberolachnus  Mordwilko  

18 

Tetraphis  Horvath 

83 

Tullgrenia  Van  der  Goot.  . 

79 

Tetrenema  Derbe  ?  

73 

Tychea  Koch.. 

71 

Thecabius  Koch 

77 

Tychea  Passermi 

79 

Thelaxes  Westwood 

21 

Tycheoides  Schouteden 

79 

Therioaphis  Walter  

28 

Unilachnus  Wilson  .  . 

17 

Thomasia  Wilson 

35 

Uraphis  Del  Guercio 

43 

Thoracaphis  Van  der  Goot  

87 

Vannna  TTeydfin 

21 

Thripsaphis  G  illette  

30 

Vesiculaphis  Del  Guercio 

51 

Todolachnus  Matsumura  

15 

Watabura  Matsumura.  .  . 

69 

Toxoptera  Koch  

51 

Wilsonia  Baker 

16 

Trama  Heyden  

19 

51 

Tranaphis  Walker  

33 

Yezfihnra  \fatsiiTpiira 

"    42 

Travaresiella  Del  Guercio  

21 

Yezocallis  Matsumura  

30 

Trichosiphum  Pergande.  .  . 

37 

Yezosiphum  Matsumura.  .  . 

49 

PLATE  I. 

A. — Anoeda  querd,  apterous  form. 
B. — Anoeda  querd,  wings. 
C. — Anoeda  querd,  antenna  of  alate  form. 
D. — Anoeda  corni,  fore  wing. 
E. — Anoeda  corni,  antenna  of  alate  form. 
F. — Anoeda  corni,  cornicle  of  alate  form. 
G. — Nippolachnus  pyri,  fore  wing. 
H. — Nippolachnus  pyri,  head  of  alate  form. 
I. — Anoeda  corni,  head  of  alate  form. 
J. — Nippolachnus  pyri,  cornicle  of  alate  form. 
K. — Nippolachnus  pyri,  antenna  of  alate  form. 
L. — Eulachnus  agilis,  body  of  alate  form. 
M. — Eulachnus  agilis,  fore  wing. 
N. — Eulachnus  agilis,  antenna  of  alate  form. 
0. — Eulachnus  agilis,  cornicle  of  alate  form. 
P. — Eulachnus  rileyi,  head  of  alate  form. 
Q. — Eulachnus  rileyi,  cornicle  of  alate  form. 
R. — Eulachnus  rileyi,  segment  III,  antenna  of  alate  form. 
S. — Essigella  californica,  body  of  alate  form. 
T. — Essigella  californica,  fore  wing. 
IT. — Essigella  californica,  antenna  of  alate  form. 
V. — Essigella  californica,  cornicle  of  alate  form. 
W. — Essigella  californica,  head  of  alate  form. 
X. — Essigella  californica,  cauda  of  alate  form. 
Y. — Essigella  californica,  antenna  of  apterous  form 
94 


Bui.  826,  U.  S.   Dept.  of  Agriculture. 


PLATE  I. 


04 


Bui.  826,   U.  S.   Dept.  of  Agriculture. 


PLATE  1 1 . 


PLATE   II. 

A. — Dilachnus  ponderosae,  fore  wing. 

B . — Dilachnus  ponderosae,  cornicle  of  apterous  form. 

C. — Dilachnus  ponderosae,  head  of  alate  form. 

D. — Schizolachnus  tomentosus,  fore  wing. 

E. —  Unilachnus  parvus,  fore  wing. 

F. —  Unilachnus  parvus,  cornicle  of  alate  form. 

G. —  Unilachnus  parvus,  head  of  alate  form. 

H. — Longistigma  caryae,  wings. 

I. — Longistigma  caryae,  apterous  form. 

J. — Longistigma  caryae,  cornicle. 

K. — Longistigma  caryae,  cornicle. 

L. — Longistigma  caryae,  antenna  of  alate  form. 

M. — Stomaphis  quercus,  alate  form. 

N. — Stomaphis  quercus,  head  of  alate  form. 

O. — Stomaphis  quercus,  cornicle  of  alate  form. 

P. — Stomaphis  quercus,  antenna  of  alate  form. 

Q. — Stomaphis  quercus,  cauda  and  anal  plate. 

R. — Stomaphis  quercus,  pits  on  rostrum. 

S. — Pterochlorus  roboris,  wing. 

T. — Pterochlorus  roboris,  head  of  alato  form. 

U. — Pterochlorus  roboris,  cornicle  of  alate  form. 

V. — Pterochlorus  riminalis,  wings. 

W. — -Pterochlorus  viminalis,  abdomen  of  alate  form. 

X. — Pterochlorus  viminalis,  head  of  alate  form. 

95 


PLATE  III. 

A. —  Thelaxes  dryophila,  apterous  form. 
B. —  Thelaxes  dryophila,  wings. 
C. — Thelaxes  dryophila,  cornicle  of  alate  form. 
D. — Thelaxes  dryophila,  cauda  of  alate  form. 
E. — Thelaxes  dryophila,  cauda  of  apterous  form. 
F. — Thelaxes  dryophila,  antenna  of  alate  form. 
G. — Glyphina  betulae,  apterous  form. 
H. — Glyphina  betulae,  wings. 
I. — Glyphina  betulae,  antenna  of  alate  form. 
J. — Glyphina  betulae,  cornicle  of  alate  form. 
K. — Glyphina  betulae,  cauda  and  anal  plate  of  alate  form. 
L. — Glyphina  betulae,  cauda  and  anal  plate  of  apterous  form. 
M. — Neotrama  delguercioi,  apterous  form. 
N. —  Trama  troglodytes,  apterous  form. 
O. — Neotrama  delguercioi,  cornicle. 
P. — Protrama  radicis,  apterous  form. 
Q. — Protrama  radicis,  fore  wing. 
R. — Protrama  radicis,  cornicle  of  alate  form. 
S. — Protrama  radicis,  antenna  of  alate  form. 
T. — Protrama  radicis,  tarsus  of  alate  form. 
96 


Bui.  826,   U.  S.   Dept.  of  Agriculture. 


PLATE  III. 


Bui.  826,   U.  S.   Dept.  of  Agriculture. 


PLATE  I V. 


PLATE  IV. 

A. — Amphorophora  rubi,  head  of  apterous  form. 

B. — Amphorophora  rubi,  cornicle. 

0. — Amphorophora  rubi,  cauda. 

D. — Rhopalosiphoninus  latysiphon,  head  of  alate  form. 

E. — Rhopalosiphoninus  latysiphon,  cornicle  of  alate  form. 

F. — Rhopalosiphoninus  latysiphon,  cauda  of  alate  form. 

G. — Myzocallis  coryli,  cornicle. 

II. — Myzocallis  coryli,  cauda  and  anal  plate. 

I. — Callipterus  juglandis,  cornicle. 

J. — Callipterus  juglandis,  cauda  and  anal  plate. 

K.—  Tlttrioaphis  tiliae,  cornicle. 

L. —  Therioaphis  tiliae,  cauda  and  anal  plate. 

M. — Monellia  caryella,  cauda  and  anal  plate. 

N. — Monellia  caryella,  cornicle. 

0. — Chromaphis  juglandicola,  cornicle. 

P. — Chromaphis  juglandicola,  cauda  and  anal  plate. 

Q. — Euceraphis  betulae,  cauda  and  anal  plate. 

R. — Euceraphis  betulae,  cornicle. 

S. — Calaphis  betulella,  cauda  and  anal  plate. 

T. — Calaphis  betulella,  cornicle. 

U. — Calaphis  betulella,  fore  wing. 

V. — Saltusaphis  scirpus,  head  of  apterous  form. 

W. — Saltusaphis  scirpus,  cauda  and  anal  plate. 

X. —  Thripsaphis  balli,  head  of  apterous  form. 

Y. — Neothomasia  populicola,  cornicle. 

Z. — Neothomasia  populicola,  cauda  and  anal  plate. 

AA. — Periphyllus  negundinis,  cauda  and  anal  plate. 

BB. — Periphyllus  negundinis,  cornicle. 

CC. — Chaitophorus  populi,  cauda  and  anal  plate. 

DD. — Symydobius  oblongus,  cornicle. 

EE. — Symydobius  oblongus,  cauda  and  anal  plat/  . 

FF. — Phyllaphis  fagi,  cauda  and  anal  plate. 

GG. — Phyllaphis  fagi,  cornicle. 

till. — Tamalea  coweni,  cornicle. 

II. — Tamalea  coweni,  cauda  and  anal  plate. 

JJ. — Drepanaphis  acerifolii,  oviparous  abdomen. 

KK. — Drepanaphis  acerifolii,  cauda  and  anal  plate  of  alate  form. 

LL. — Drepanaphis  acerifolii,  cornicle. 

MM. — Drepanosiphum  platanoides,  cornicle. 

NN. — Melanoxantherium  populifoliae,  cornicle. 

00. — Melanoxantherium  populifoliae,  cauda  and  anal  plate. 

PP. — Pterocomma  populeus,  cornicle. 

141613°— 20— Bull.  826 7  97 


PLATE  V. 

A. — Eutrichosiphum  pasaniae,  antenna  of  alate  form. 

13. — Eutrichosiphum  pasaniae,  wings. 

C. — Eutrichosiphum  pasaniae,  cornicle  of  apteroua  form. 

D. — Eutrochosiphum  pasaniae,  cornicle  of  alate  form. 

E. — Eutrichosiphum  pasaniae,  antenna  of  apterous  form. 

F. — Greenidea  anonae,  apterous  form. 

G. — Greenidea  artocarpi,  wings. 

H. — Greenidea  artocarpi,  cauda  of  apterous  form. 

I. — Greenidea  artocarpi,  cornicle  of  apterous  form. 

J. — Greenidea  anonae,  abdomen  of  alate  form. 

K. — Greenidea  anonae,  head  of  alate  form. 

L. — Greenideoida  elongata,  wings. 

M. — Greenideoida  sp. ,  head  of  alate  form. 

N. — Greenideoida  sp.,  cornicle  of  alate  form. 

O. — Greenideoida  sp.,  third  antennal  segment  of  alate  form?} 

P. — Greenideoida  hannae,  cornicle  of  apterous  form. 

Q. — Setaphis  luteus,  caudal  portion  of  apteroua  form. 

R. — Setaphis  luteus,  wings. 

S. — Setaphis  luteus,  head  of  alate  form. 

T. — Setaphis  luteus,  cornicle  of  alate  form. 

IT. — Setaphis  luteus,  spine  of  alate  form. 

V. — Setaphis  luteus,  cornicle  of  apterous  form. 

W. — Setaphis  luteus,  antenna  of  alate  form. 

X. — Setaphis  luteus,  antenna  of  apterous  form. 

98 


Bui.  826,   U.  S.   Dept.  of  Agriculture. 


PLATE  V. 


rv 


UJ 

O 


Bui.  826.  U.  S.  Dept.  of  Agriculture. 


PLATE  VI. 


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^*.(     -          V.  :  -'  .'•  "-•-•X  .%-  ./.;•-*—. 


PLATE  VI. 

A. — Acandus  lychnidis,  cornicle  of  alate  form. 

B. — Acaudus  lychnidis,  cauda  of  alate  form. 

C. — Anuraphis  amygdali,  cornicle  of  alate  form. 

D. — Anuraphis  amygdali,  cauda  of  alate  form. 

E. — Anuraphis  amygdali,  head  of  alate  form. 

F. — Anuraphis  carotae,  cornicle  and  cauda  of  alate  form. 

G. — Aphis  sambuci,  head  of  alate  form. 

H. — Aphis  sambuci,  cornicle  of  alate  form. 

I. — Aphis  sambuci,  cauda  of  alate  form. 

J. — Brevicoryne  brassicae,  cauda  of  alate  form. 

K. — Brevicoryne  brassicae,  cornicle  of  alate  form. 

L. — Aspidaphis  polygonii,  cornicle  of  alate  form. 

M. — Aspidaphis  polygonii,  cornicle  more  enlarged. 

N. — Aspidaphis  polygonii,  cauda  of  apterous  form. 

0. — Aspidaphis  polygonii,  head  of  apterous  form. 

P. — Atarsos  grindeliae,  cornicle  of  alate  form. 

Q. — Atarsos  grindeliae,  head  of  alate  form. 

R. — Atarsos  grindeliae,  cauda  of  alate  form. 

S. — Atarsos  grindeliae,  tibia  of  alate  form. 

T. — Brachycolus  stellariae,  cornicle  of  alate  form. 

U. — Brachycolus  stellariae,  cauda  of  alate  form. 

V. — Carolinaia  cyperi,  cornicle  of  alate  form. 

W. — Carolinaia  cyperi,  cauda  of  alate  form. 

X. — Cavariella  pastinacae,  cornicle  of  alate  form. 

Y. — Cavariella  pastinacae,  cauda  of  alate  form. 

Z. — Cavariella  pastinacae,  tubercle  of  alate  form. 

AA. — Hyadaphis  xylostei,  cauda  of  apterous  form. 

BB. — Hyadaphis  xylostei,  cornicle  of  apterous  form. 

CC. — Toxoptera  aurantiae,  head  of  alate  form. 

DD. —  Toxoptera  aurantiae,  cornicle  of  alate  form. 

EE. —  Toxoptera  aurantiae,  cauda  of  alate  form. 

FF. — Rhopalosiphum  rufomaculata,  cornicle  of  alate  form. 

GG. — Rhopalosiphum  rufomaculata,  head  of  alate  form. 

HH. — Rhopalosiphum  rufomaculata,  cauda  of  alate  form. 

II. — Rhopalosiphum  nymphaeae,  cornicle  of  alate  form. 

JJ. — Rhopalosiphum  nymphaeae,  cauda  of  alate  form. 

KK. — Pergandeidia  ononidis,  cauda  and  cornicle  of  alate  form. 

LL. — Pergandeidia  trirhodus,  cauda  of  apterous  form. 

MM. — Pergandeidia  trirhodus,  cornicle  of  apterous  form. 

NN. — Liosomaphis  berberidis,  cornicle  and  cauda  of  apterous  form. 

00. — Liosomaphis  berberidis,  cauda  of  alate  form. 

PP. — Cryptosiphum  artemesuie,  cornicle  of  apterous  form. 

QQ. — Cryptosiphum  artemesiae,  cauda  of  apterous  form. 

RR. — Hyalopterus  arundinis,  cornicle  of  alate  form. 

SS. — Hyalopterus  arundinis,  cauda  of  alate  form. 

TT. — Hyalopterus  deformans,  cauda  of  apterous  form. 

UU. — Hyalopterus  deformans,  cornicle  of  apterous  form. 

W. — Hyalopterus  deformans,  cauda  of  alate  form. 

WW. — Hyalopterus  deformans,  cornicle  of  alate  form. 

99 


PLATE  VII. 

A. — Siphonatrophia  cupressi,  apterous  form. 
B. — Siphonatrophia  cupressi,  wings. 
C. — Siphonatrophia  cupressi,  antenna  of  alate  form. 
D. — Siphonatrophia  cupressi,  cauda  of  alate  form. 
E. — Siphonatrophia  cupressi,  antenna  of  apterous  form. 
F. — Sanbornia  juniperi,  apterous  form. 
G. — Sanbornia  juniperi,  \vings. 
H. — Sanbornia  juniperi,  antenna  of  alate  form. 
I. — Sanbornia  juniperi,  head  of  apterous  form  from  beneath. 
J. — Sanbornia  juniperi,  cauda  of  alate  form. 
K. — Sanbornia  juniperi,  cornicle. 
L. — Sanbornia  juniperi,  tarsus. 
M. —  Vesiculaphis  caricis,  apterous  form. 
N. —  Vesiculaphis  caricis,  cornicle  of  alate  form. 
0. —  Vesiculaphis  caricis,  fore  wing. 
P. —  Vesiculaphis  caricis,  antenna  of  alate  form. 
Q. —  Vesiculaphis  caricis,  cauda  of  alate  form. 
R. — Acanthaphis  rubi,  apterous  form. 
S. — Acanthaphis  rubi,  head  of  apterous  form. 
T. — Acanthaphis  rubi,  cornicle  of  apterous  form. 
U. — Acanthaphis  rubi,  antenna  of  apterous  form. 
100 


Bui.  826,   U.  S.   Dept.  of  Agriculture. 


PLATE  VI  I. 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  VIM. 


PLATE  VIII. 

A. — Capitophorus  shepardiae,  cornicle  of  alate  form. 

B. — Capitophorus  shepardiae,  head  of  alate  form. 

0. — Capitophorus  shepardiae,  cauda  of  alate  form. 

D. — Anomalaphis  comperi,  apterous  form. 

E. — Anomalaphis  comperi,  antenna  of  apterous  form. 

F. — Anomalaphis  comperi,  wings. 

G. — Cervaphis  schoutedeniae,  extremity  of  abdomen. 

H. — Illinoia  liriodendri,  head  of  alate  form. 

I. — Illinoia  liriodendri,  cornicle  of  alate  form. 

J. — Illinoia  liriodendri,  cauda  of  alate  form. 

K. — Myzus  cerasi,  head  of  apterous  form. 

L. — Myzus  cerasi,  cauda  of  apterous  form. 

M. — Myzus  cerasi,  cornicle  of  alate  form. 

N. — Phorodon  humuli,  head  of  apterous  form. 

0. — Phorodon  humuli,  cauda  of  apterous  form. 

P. — Phorodon  humuli,  cornicle  of  apterous  form. 

Q. — Phorodon  humuli,  head  of  alate  form. 

R. — Macrosiphonella  sanborni,  head  of  alate  form. 

S. — Macrosiphonella  sanborni,  cornicle  of  alate  form. 

T. — Macrosiphonella  sanborni,  cauda  of  alate  form. 

IT. — Macrosiphum  rosae,  head  of  alate  form. 

V. — Macrosiphum  rosae,  cauda  of  alate  form. 

W. — Macrosiphum  rosae,  cornicle  of  alate  form. 

X.— Myzus  persicae,  head  of  apterous  form. 

Y. — Myzus  persicae,  cornicle  of  alate  form. 

Z. — Myzus  persicae,  cauda  of  alate  form. 

AA. — Microparsus  variabilis,  cauda  of  alate  form. 

BB. — Microparsus  variabilis,  wings. 

CO. — Microparsus  variabilis,  cornicle  of  alate  form. 

DD. —  Microparsus  variabilis,  head  of  alate  form. 

EE. — Idiopterus  nephrolepidis ,  cornicle  of  alate  form. 

FF. — Idiopterus  nephrolepidis,  fore  wing. 

GG. — Idiopterus  nephrolepidis,  head  of  alate  form. 

HH. — Idiopterus  nephrolepidis,  cauda  of  alate  form. 

II. — Pentalonia  nigronervosa,  head  of  apterous  form. 

JJ. — Pentalonia  nigronervosa,  wings. 

KK. — Pentalonia  nigronervosa,  cauda  of  alate  form. 

LL. — Pentalonia  nigronervosa,  head  of  alate  form. 

MM. — Pentalonia  nigronervosa,  cornicle  of  alate  form. 

101 


PLATB  IX. 

A. — Mindarus  abietinus,  winga. 
B. — Mindarus  abietinus,  fore  wing  showing  tracheae. 
C. — Mindarus  abietinus,  cauda  of  alate  form. 
D. — Mindarus  abietinus,  antenna  of  alate  form. 
E. — Mindarus  abietinus,  abdomen  of  pupa. 
F. — Mindarus  abietinus,  cornicle  of  alate  form. 
G. — Colopha  ulmicola,  antenna  of  alate  form. 
H. — Colopha  ulmicola,  antenna  of  stem  mother. 
I. — Colopha  ulmicola,  wings. 
J. — Colopha  ulmicola,  apterous  form. 
K. — Colopha  ulmicola,  antenna  of  apterous  form. 
L. — Colopha  ulmicola,  antenna  of  apterous  form. 
M. — Eriosoma  lanigerum,  apterous  form. 
N. — Eriosoma  lanigerum,  wax  plate. 
0. — Eriosoma  lanigerum,  wax  reservoir. 
P. — Eriosoma  lanigerum,  fore  wing. 
Q. — Eriosoma  lanigerum,  wing  pad  showing  trachea. 
R. — Eriosoma  lanigerum,  cornicle. 
S. — Eriosoma  lanigerum,  antenna  of  alate  form. 
T. — Eriosoma  lanigerum,  oviparous  female,  showing  egg. 
U. — Georgia  ulmi,  antenna  of  alate  form. 
V. — Georgia  ulmi,  wings. 

W. — Georgia  ulmi,  distal  segment  of  antenna  of  alate  form. 
X. — Georgia  ulmi,  cornicle  of  alate  form. 
Y. — Georgia  ulmi,  cauda  and  anal  plate  of  alate  form. 
Z. — Georgia  ulmi,  head  of  alate  form. 
102 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  IX. 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  X. 


o 


PLATE  X. 

A. — Gobaishia  pallida,  antenna  of  alate  form. 

B. — Gobaishia  pallida,  antenna  of  stem  mother. 

C. — Gobaishia  pallida,  wings. 

B. — Gobaishia  pallida,  abdomen  of  alate  form  showing  cornicle. 

E. — Gobaishia  pallida,  fore  wing. 

F. — Gobaishia  pallida,  abdomen  of  pupa. 

G. — Gobaishia  pallida,  distal  segment  of  antenna  of  alate  form  from  above. 

H. —  Tetraneura  ulmifoliae,  antenna  of  alate  form. 

I. —  Tetraneura  ulmifoliae,  antenna  of  stem  mother. 

J. —  Tetraneura  ulmifoliae,  antenna  of  apterous  form. 

K. —  Tetraneura  ulmifoliae,  wings. 

L. —  Tetraneura  ulmifoliae,  wings  showing  tracheae. 

M. —  Tetraneura  ulmifoliae,  cauda  and  anal  plate. 

N. — Cornaphis  populi,  stem  mother. 

0. — Cornaphis  populi,  antenna  of  stem  mother. 

P. — Cornaj>his  populi,  wings. 

Q. — Cornaphis  populi,  cornicle  of  alate  form. 

R. — Cornaphis  populi,  antenna  of  alate  form. 

S. — Cornaphis  populi,  fore  wing. 

T. — Cornaphis  populi,  head  showing  horn. 

U. — Dryopeia  bella,  apterous  form. 

V. — Dryopeia  bella,  wings. 

W. — Dryopeia  bella,  antenna  of  apterous  form. 

X. — Dryopeia  bella,  antenna  of  alate  form. 

Y. — Dryopeia  bella,  cornicle  of  apterous  form. 

108 


PLATE   XI. 

A. — Mordwilkoja  vagabunda,  stem  mother. 
B. — Mordwilkoja  vagabunda,  antenna  of  stem  mother. 
C. — Mordwilkoja  vagabunda,  antenna  of  alate  form. 
D. — Mordwilkoja  vagabunda,  wings. 
E. — Mordwilkoja  vagabunda,  thorax  of  alate  form. 
F. — 'Mordwilkoja  vagabunda,  cauda  of  alate  form. 
G. — Mordwilkoja  vagabunda,  cornicle  of  alate  form. 
H.— Asiphum  sp.  stem  mother. 
I. — Asiphum  sp.  antenna  of  stem  mother. 
J. — Pachypappella  vesicalis,  wings. 

K. — Pachypappella  vesicalis,  cornicles  and  cauda  of  alate  form. 
L. — Pachypappella  vesicalis,  antenna  of  alate  form. 
M. — Pachypappella  vesicalis,  wax  plate. 
N. — Pemphigus  bursarius,  antenna  of  stem  mother. 
O. — Pemphigus  bursarius,  wings. 
P. — Pemphigus  bursarius,  antenna  of  alate  form. 
Q. — Pemphigus  bursarius,  antenna  of  alate  form. 
~R.~Pemphigus  bursarius,  antenna  of  stem  mother. 
S. — Pemphigus  bursarius,  wax  plate. 
T. — Pemphigus  bursarius,  cornicle  of  alate  form. 
U. — Pemphigus  populicaulis,  antenna  of  stem  mother. 
V. — Gobaishia  ulmifuscus,  antenna  of  stem  mother. 
W. — Phloeomyzus  passerini,  apterous  form. 
X. — Phloeomyzus  passerini,  antenna  of  apterous  form. 
Y. — Phloeomyzus  passerini,  wings. 
Z. — Phloeomyzus  passerini,  antenna  of  apterous  form. 
AA. — Phloeomyzus  passerini,  antenna  of  alate  form. 
BB. — Phloeomyzus  passerini,  wax  plate. 
104 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  XI. 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  XI  I. 


o 


CO 


PLATE   XII. 

A.— Aploneura  lentici,  apterous  form. 

~B.~Aploneura  lentici,  antenna  of  apterous  form. 

C. — Aploneura  lentici,  antenna  of  alate  form. 

D. — Aploneura  lentici,  wings. 

E.—  Aploneura  lentici,  abdominal  wax  plate. 

F. — Melaphis  rhois,  antenna  of  apterous  form. 

G. — Melaphis  rhois,  antenna  of  alate  form. 

H. — -Melaphis  rhois,  wings. 

I. — Melaphis  rhois,  abdomen  of  alate  form. 

J. — -Melaphis  rhois,  antenna  of  alate  form. 

K. — Melaphis  chinensis,  fore  wing. 

L. — Nurudea  ibofushi,  wings. 

M. — Nurudea  ibofushi,  antenna  of  alate  form. 

N. — -Nurudea  shiraii,  wings. 

O. — Nurudea  shiraii,  antenna  of  alate  form. 

P. — Nurudea  rosea,  fore  wing. 

Q. — Nurudea  rosea,  antenna  of  alate  form. 

R.—  Asiphum  tremulae,  stem  mother. 

S. — Asiphum  tremulae,  antenna  of  stem  mother. 

T. — Asiphum  tremulae,  antenna  of  alate  form. 

U. — Asiphum  tremulae,  wings. 

V. — Asiphum  tremulae,  abdomen  of  alate  form. 

W. — Asiphum  tremulae,  wax  plates. 

X. — Asiphum  tremulae,  antenna  of  apterous  form. 

105 


PLATE  XIII. 

A. — Neoprociphilus  attenuatus,  forewing. 
B. — Neoprociphilus  attenuatus,  antenna  of  alate  form. 
C. — Neoprociphilus  attenuatus,  segment  V  of  antenna  of  alate  form. 
D.— Neoprociphilus  attenuatus,  antenna  of  apterous  form. 
E. — Neoprociphilus  attenuatus,  antenna  of  male. 
F. — Neoprociphilus  attenuatus,  antenna  of  oviparous  female. 
G. — Prociphilus  bumeliae,  antenna  of  alate  female. 
H. — Prociphilus  bumeliae,  antenna  of  stem  mother. 
I. — Prociphilus  bumeliae,  wings. 
J. — Prociphilus  bumeliae,  abdomen  of  alate  form. 
K. — Prociphilus  bumeliae,  thoracic  wax  plates. 
L. — Prociphilus  bumeliae,  head  wax  plates. 
M. — Prociphilus  xylostei,  thoracic  wax  plates. 
N. — Prociphilus  bumeliae  (poscheringi) ,  thoracic  wax  plates. 
0. — Thecabius  affinis,  antenna  of  stem  mother. 
P. —  Thecabius  affinis,  antenna  of  alate  form. 
Q. — Thecabius  affinis,  wings. 
R. — Thecabius  affinis,  thorax  of  alate  form. 
S. — Thecabius  affinis,  head  wax  plates  of  stem  mother. 
T. — Thecabius  affinis,  abdominal  wax  plate. 
U. — Thecabius  affinis,  abdominal  wax  plates  of  stem  mother. 
V. — Forda  sp.,  apterous  form. 
W. — Forda  olivacea,  antenna  of  alate  form. 
X. — Forda  trivialis,  antenna  of  alate  form. 
Y. — Forda  marginata,  antenna  of  alate  form. 
Z. — Forda  olivacea,  fore  wing. 
AA. — Forda  for  micaria,  antenna  of  apterous  form. 
106 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  XII I. 


o 


Bui.  826.  U.  S.  Dept.  of  Agriculture. 


PLATE  XIV. 


PLATE  XIV. 

A. — Geoica  squamosa,  apterous  form. 

B. — Geoica  squamosa,  antenna  of  alate  form. 

C. — Geoica  squamosa,  eye  of  apterous  form. 

D. — Geoica  squamosa,  wings. 

E. — Geoica  squamosa,  caudal  extremity  of  apterous  form. 

F. — Geoica  squamosa,  squama. 

G. — Geoica  squamosa,  four-segmented  antenna  of  apterons  form. 

II. — Geoica  phaseoli,  antenna  of  alate  form. 

I.— Geoica  phaseoli,  antenna  of  apterous  form. 

J. — Geoica  radicicola,  antenna  of  alate  form. 

K. — Geoica  lucifuga,  antenna  of  alate  form. 

1^.— Paracletus  cimiciformis,  apterous  form. 

M. — Paracletus  cimiciformis,  eye  of  apterous  form. 

N. — Paracletus  cimiciformis,  intermediate  eye. 

O. — Paracletus  cimiciformis,  median  thoracic  wax  plate  of  alate  form. 

P. — Paracletus  cimiciformis,  foot  of  alate  form  with  one  claw. 

Q. — Paracletus  cimiciformis,  ?     enna  of  apterous  form. 

R. — Paracletus  cimiciformis,  .Brewing. 

S. — Paracletus  cimiciformis,  antenna  of  alate  form. 

T. — Uamamelistes  spinosus,  hibernating  apterons  form. 

U. — ITamamelistes  spinosus,  wings. 

V. — Uamamelistes  spinosus,  cauda  and  anal  plate  of  alate  form. 

W. — Uamamelistes  spinosus,  antenna  of  alate  form. 

X. — Hamamelistes  spinosus,  portion  of  same,  more  enlarged. 

Y. — ITormaphis  Jiamamelidis,  wings. 

Z. — Hormaphis  Jiamamelidis,  antenna  of  alate  form. 

AA. — Tlormaphis  Jiamamelidis,  antenna  of  stem  mother. 

BB. — TTormaphis  Jiamamelidis,  apterous  form. 

CC.—TTormaphis  Jiamamelidis,  antenna  of  apterous  form. 

DD. — Tlormaphis  hamamelidis,  cauda  and  anal  plate  of  apterous  form. 

EE. — ITormaphis    hamamelidis,  dorsal  wax  pores. 

FF. — TTormaphis  hamamelidis,  cauda  and  anal  plate  of  alate  form. 

107 


PLATE  XV. 

A. — Oregma  lanigera,  apterous  form. 

B. — Oregma  lanigera,  antenna  of  alate  form. 

C. — Oregma  lanigera,  cauda  and  anal  plate  of  alate  form. 

D. — Oregma  lanigera,  wings. 

E — Oregma  lanigera,  end  of  abdomen  of  apterous  form. 

F. — Oregma  lanigera,  antenna  of  apterous  form. 

G. — Oregma  bambusae,  head  of  apterous  form. 

H. — Glyphinaphis  bambusae,  apterous  form. 

I. — Glyphinaphis  bambusae,  antenna  of  apterous  form. 

j. — Glyphinaphis  bambusae,  cornicle  of  apterous  form. 
K.— Glyphinaphis  bambusae,  cauda  and  anal  plate. 

L. — Mansakia  miyabei,  wings. 

M. — Ceratoglyphina  bambusae,  apterous  form. 

X.— Ceratoglyphina  bambusae,  cauda  and  anal  plate. 

0. — Ceratoglyphina  bambusae,  cornicle  of  apterous  form. 

P.— Ceratoglyphina  bambusae,  antenna  of  apterous  form. 

Q. — Astegopteryx  distychii,  antenna  of  alate  form. 

R. — Astegopteryx  distychii,  fore  wing. 

S.— Astegopteryx  distychii,  anal  plate  of  alate  form. 

T. — Astegopteryx  distychii,  cauda  of  alate  form. 

U. — Astegopteryx  distychii,  end  of  abdomen. 

V. — Astegopteryx  styracophila,  fore  wing. 
W. — Astegopteryx  gallarum,  fore  wing. 

"S..— Astegopteryx  gallarum,  tip  of  antenna  of  alate  form. 
108 


Bui.  826.  U.  S.  Dept.  of  Agriculture. 


PLATE  XV. 


o 


OQ  = 


Bui.  826,  U.  S.  Dept.  of  Agriculture. 


PLATE  XVI. 


PLATE  XVI. 

A. — Aleurodaphis  blumeac,  apterous  form. 
B. — Aleurodaphis  blumeae,  antenna  of  apterous  form. 
C. — Aleurodaphis  blumeae,  dorsal  wax  pores. 
D. — Aleurodaphis  blumeae,  margin. 
E. — Aleurodaphis  blumeae,  cauda  and  anal  plate. 
F. — Cerataphis  lataniae,  apterous  form. 
G. — Cerataphis  lataniae,  wings. 

H.— Cerataphis  lataniae,  cauda  and  anal  plate  of  apterous  form. 
I.  —  Cerataphis  lataniae,  antenna  of  apterous  form. 
J. — Cerataphis  lataniae,  antenna  of  alate  form. 
K. — Cerataphis  lataniae,  anal  plate  of  alate  form. 
L. — Cerataphis  lataniae,  cauda  of  alate  form. 
M. — Cerataphis  lataniae,  head  of  apterous  form,  central  view. 
N. — Thoracaphis  arboris,  apterous  form. 
O. — Thoracaphis  arboris,  antenna  of  apterous  form. 
P. — Thoracaphis  arboris,  cauda  and  anal  plate  of  apterous  form. 
Q. —  Thoracaphis  ficus,  antenna  of  apterous  form. 
R. — Thoracaphis  ficus,  extremity  of  abdomen,  apterous  form. 
S. — Thoracaphis  castaneae,  wings. 

T. — -Thoracaphis  castaneae,  distal  segment  of  antenna  of  alate  form. 
U. — Thoracaphis  castaneae,  cauda  and  anal  plate  of  alate  form. 
V. — Thoracaphis  castaneae,  cornicle  of  alate  form. 
W. —  Thoracaphis  castaneae,  antenna  of  alate  form. 

109 


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